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GEOLOGY Vol. 27 No. i 

LONDON: 1976 






Pp. 1-30 ; 3 Plates ; 7 Text-figures 



GEOLOGY Vol. 27 No. i 

LONDON: 1976 


(natural history), instituted in 1949, is 
issued in five series corresponding to the Scientific 
Departments of the Museum, and an Historical series. 

Parts will appear at irregular intervals as they 
become ready. Volumes will contain about three or 
four hundred pages, and will not necessarily be 
completed within one calendar year. 

In 1965 a separate supplementary series of longer 
papers was instituted, numbered serially for each 

This paper is Vol. 27, No. i, of the Geological 
iPalaeontological) series. The abbreviated titles of 
periodicals cited follow those of the World List of 
Scientific Periodicals. 

World List abbreviation : 
Bull. Br. Mus. nat. Hist. (Geol. 

ISSN 0007- 1471 

Trustees of the British Museum (Natural History), 1976 


Issued 4 February, 1976 Price £2.85 





Abstract 3 


Introduction .... 



Description .... 

a. Skull morphology . 

b. Postcranial elements 

c. Measurements 






Comparisons with Recent material 



Discussion and conclusions 



Systematic description 






References .... 






Prophaethon shruhsolei Andrews of the Lower Eocene, known from an imperfect skeleton, has 
been previously assigned to the Phaethontidae within the Pelecaniformes. After further 
preparation the specimen is redescribed. In comparison with Recent species of Pelecani- 
formes, Charadriiformes and Procellariiformes it was found to share some characters with all 
three, and could not with any certainty be assigned to any one of them. It is proposed, there- 
fore, that Prophaethon should be considered as a representative of a new monotypic order, 
Prophaethontiformes, and it is suggested that it represents an early link between these Recent 
orders. This hypothesis is discussed and a diagnosis of the new order is given. 


In 1899 C. W. Andrews described a fossil bird which W. H. Shrubsole had collected 
from the London Clay of the Isle of Sheppey, Kent, and had presented to the British 
Museum (Natural History). When found, the specimen consisted of a clay nodule 
in which one side of the skull and limb fragments were exposed. It was partially 
prepared at the Museum and when described and figured it showed the entire dorsal, 
posterior and right lateral views of the skull with the lower mandible in place. The 
orbit was also partly cleared of matrix back to the quadrate and down to the quad- 
rat ojugal bar. The left side of the pelvis, which was somewhat obscured by the 
overlying skull, was also prepared, and a femur and the anterior portion of a broken 
tibiotarsus were laterally exposed. 

From the general appearance of the skull Andrews concluded that it belonged to a 
pelecaniform bird. In the subsequent discussion of its affinities only pelecaniform 
families were referred to, and one may conclude that once he had decided on its 
possible ordinal status he did not compare it with material other than that within 
this taxon. 


He compared the specimen systematically with examples of the various pelecani- 
form families and concluded that it most nearly resembled the genus Phaethon which 
constitutes the family Phaethontidae. In spite of some differences he thought that 
it was ancestral to Phaethon and named it Prophaethon shrubsolei, regarding the 
general similarities of the skull and deep fronto-nasal hinge with a prominent 
frontal ridge as evidence of affinity. 

He noted that the nostrils of Prophaethon, unlike the holorhinal ones of Phaethon, 
approximated to the schizorhinal condition, but quoted Pycraft (1898) to the effect 
that on the skull of the young of Phaethon the nostrils show a nearly schizorhinal 
condition. He also noted that the pelvis was narrow and more closely resembled 
that of Sida and Phalacrocorax. He regarded the nostril condition as ancestral and 
concluded that the narrow pelvis indicated that, unlike the Recent Phaethon, 
Prophaethon was probably a good swimmer and diver, and that the extent to which 
Phaethon had diverged from this condition was a result of post-Eocene evolution. 

During work towards a comprehensive review of the British Lower Eocene birds, 
still in progress, we decided that the specimen was suitable for further preparation 
by modern techniques. This was undertaken by Mr F. M. P. Howie of the Palae- 
ontological Laboratory of the Museum. X-ray photographs were used to trace the 
position and extent of the bones present, and the specimen was then carefully 
prepared. As a result of this work most of the elements have now been separated 
and cleaned. The skull can be examined in all aspects including the palate (the 
lack of access to which Andrews regretted). The lower jaw is now separate and 
complete. A previously unsuspected portion of sternum has been revealed, and the 
hidden part of the proximal end of the tibiotarsus is now free. The pelvis has been 
extensively cleared, but still has attached to it the proximal half of the femur, and 
also portions of the ribs. An almost complete coracoid and the blade of a scapula 
are also present, and there are fragments of vertebrae and broken portions of limb- 
bones partly embedded in more resistant matrix. 

In the following sections the various parts have been described. The terminology 
follows that of Jollie (1957) for the skull, and that of Howard (1929) for the post- 
cranial skeleton. From the characters now available the species does not appear to 
be closely allied to Phaethon and some characters are apparently shared with non- 
pelecaniform taxa. 


a. Skull morphology 

Cranial and orbital structures. Viewed dorsally (Fig. i ; PL i, fig. A) and 
laterally (Fig. 2 ; PI. i, figs C & D) the skull as a whole shows a fairly even taper 
from its widest part in the temporal region to its narrow termination at the tip of the 
rostrum. The frontal rises a little from the dorsal edge of the parietals, with a slight 
inflation on either side in the area above the brain. From near the posterior edge 
of the orbits the frontal maintains a fairly even width anteriorly, with a shallow 
median hollow in the interorbital region, and then expands gradually at the anterior 


Fig. I . Dorsal view of skull, x i . Abbreviations : boc, basioccipital condyle ; bps, basi- 
parasphenoid ; dps, gland depression, ? salt ; exn, external nares ; fm, foramen magnum ; 
fnh, fronto-nasal hinge ; ios, interorbital septum ; mx, maxilla ; ns, nasal strut ; pes, 
prefrontal attachment area ; pi, palatine ; pmx, premaxilla ; pop, postorbital process ; 
q, quadrate ; rps, rostroparasphenoid ; tf, temporal fossa ; z, zygoma. 

end before terminating abruptly in a transverse, rounded and prominent brow 
ridge just above the fronto-nasal hinge. From the more complete left side of the 
frontal end of the specimen it would appear to be thick anteriorly, but above the 
orbits the frontal becomes very thin and its outer edge may have been irregular. 

The posterior dorsal edge of the orbit curves outwards and terminates in a promi- 
nent, posteriorly curved postorbital process, which projects at a level only a little 
below the dorsal surface of the frontal. Posterior to this process is a deep temporal 
fossa. Viewed from above (Fig. i ; PI. i, fig. A), the fossa is rounded and partly 
enclosed by the postorbital process. In spite of the depth of the fossa the internal 
margin does not extend very far onto the dorsal surface of the cranium and there is 
only a relatively small and poorly-defined depression around its inner edge, while 


Fig. 2. Left lateral view of skull, x i. Abbreviations as in Fig. i. 


posteriorly the main fossa slopes back to the line of the fronto-parietal junction. 
The fronto-parietal edge is superficially eroded and the original temporal fossa may 
have been more clearly defined. 

Posterior to the fossa the squamosal and parietal form an arch over the auditory 
meatus with a slight peak marking the posterior edge of the fossa ; the quadrate 
articulates immediately below it. On the left side the outer edges of the bones are 
damaged, while on the right side both the postorbital process and the region above 
the quadrate are broken. 

The cranium is rather small and projects forward medially with the result that, 
within the orbit, the posterior part of the frontal which forms the anterior ventral 
wall of the cranial cavity slants away postero-laterally, and also at an angle of 
c. 45° postero-ventrally. 

In the upper region of the orbit the frontal is pierced, on either side of the inter- 
orbital septum, by a large oval fenestra opening into the cranial cavity. At the 
lower posterior edge of the frontal (or possibly in the orbitosphenoid) there is a large 
optic foramen, piercing the interorbital septum and extending laterally on either 
side of it. The septum itself is fragmentary and shows an extensive irregular 
central fenestra and various small subsidiary foramina. There are grooves, pre- 
sumably to accommodate nerves, along the dorsal and ventral edges of the septum. 
The interorbital region of the frontal is thick, the roof of the orbit sloping from the 
orbit edge towards the septum. 

At its anterior dorsal end the interorbital septum is pierced by a small, vertical 
fenestra which extends up into a hollow in the ventral surface of the frontal at this 
point. Just posterior to this fenestra the septum edge forms a thickened ridge 
descending from the orbit roof and curving forward below the fenestra, becoming 
concealed by the matrix which encloses the inner nasal region. The interorbital 
fenestra and hollow may have accommodated some structure with an olfactory 
function. This area is more complete on the left side of the specimen. 

On this side the roof of the orbit shows a peculiar feature. Within the orbit, 
along its dorsal edge, there is an elongated oval depression, extending slightly 
inwards and terminating internally at an abrupt ridge where the frontal resumes its 
normal thickness. It is not clear to what extent the edge of the orbit may have been 
damaged, but there has been a reduction in thickness of the orbit roof, towards its 
edge, to accommodate a structure which was elongated, broader towards its middle, 
and dorso-ventrally flattened to an even thickness. 

At the anterior end of the orbit the frontal is thick and abruptly flattened on the 
lateral surface. In some of the Recent Pelecaniformes this condition is associated 
with the presence of a closely attached but unfused prefrontal, which may fall away 
to reveal a similar surface of attachment. It therefore seems likely that a pro- 
jecting prefrontal was originally present in Prophaethon, but has been lost. On the 
left side, within the orbit, there is an incomplete and laterally projecting flange of 
bone at the middle of the anterior edge of the interorbital septum. This might be 
the remains of the lateral ethmoid plate associated with it. In either case there is 
evidence for the existence of some structure at the anterior end of the orbital cavity, 
of a type normally associated with a prefrontal. 


Fig. 3. Ventral view of skull with left zygoma removed, x i. Abbreviations as in Fig. i. 

The jugal and quadrat ojugal form a narrow even strut of bone arising a little 
dorsal to the posterior tip of the maxillary and extending in a straight line back to 
the quadrate, continuing the line of the tomium of the upper mandible. The bar 
has been fragmented but appears to be laterally flattened along its entire length 
and shows no torsion. 

Ventrally (Fig. 3 ; PL i, fig. B), the basiparasphenoid is a small and triangular 
plate, tapering anteriorly to a smooth rostroparasphenoid and at the external 
posterior corners curving ventrally to form a pair of blunt projections only a little 
anterior to the occipital condyle. The area is damaged and pitted in places and it is 
not possible to be certain of the structure of the anterior part of the basiparasphenoid 

The posterior surface (Fig. 4 ; PI. i, fig. E) of the cranium is rather flattened, but 
a little inflated in the region of the supraoccipital. The foramen magnum is large 
and rounded and tilted slightly downwards. The parietals form a broad arch over 
the foramen. They are slightly hollowed dorsally towards the inner end, and the 


Fig. 4. Posterior view of skull, x i. Abbreviations as in Fig. i. 


outer ventral edges are prominent and rounded posteriorly. The surfaces of the 
exoccipitals are concave. They have a slightly anterior ventral slant and begin to 
taper ventrally before expanding again at the line of fusion with the posterior edge 
of the basiparasphenoid plate. 

Quadrate. In Prophaethon the right quadrate is in the normal position with the 
shaft vertical, but it has been damaged along its external side. The left quadrate 
is also in position, but the posterior end of the pterygoid has become displaced 
upwards, allowing the quadrate to tilt inwards at its lower end ; it is held in this 
position by matrix on the internal side. The tilt has displaced the dorsal articula- 
ting surface and allows a better view of this end. 

The shaft is postero-anteriorly flattened, the narrow waist at the level of the 
orbital process being about three times as broad as it is thick, and widening towards 
the dorsal and ventral ends. The dorsal head is laterally broad, the external 
flange of the otic process projecting laterally as far as does the quadrat ojugal socket 
on the ventral head. The otic process slants laterally from the waist to form a 
rather flattened process, with a shallow ridge from the base of the orbital process 
crossing the anterior surface, but with some of the posterior surface and tip broken 
away. The internal facet of the dorsal head appears to terminate as a rounded 
structure projecting internally only a little further than the inner edge of the shaft. 

The orbital process is blade-like, with an almost horizontal upper edge and a 
slightly rounded tip. From this tip it curves down towards the lower part of the 
shaft. It appears to project anteriorly at an angle to the quadrate shaft of about 70°. 

The ventral head of the quadrate is elongated laterally. The quadratojugal 
socket projects laterally and curves forwards so that the entrance of the socket is 
more anterior than the lateral alignment. The posterior side of the head curves 
back from the articulation socket to form a projecting posterior flange ; from the 
posterior side of the broken right quadrate of the specimen it would appear that 
there is a fairly deep but small groove on the lower end of the mid-shaft, cutting into 
the flange. The ventral surface of the quadrate then curves anteriorly again, but 
on the specimen the mandibular articulation surface of this region is damaged. 

Palate (Fig. 3 ; PL i, fig. B). The skull presents an example of a typical schizo- 
gnathous palate. The maxillae are broadest posteriorly at the point where they fuse 
with the palatines, and posteriorly they then taper rapidly on the external side to the 
articulation with the jugal. Anteriorly, they taper a little and extend for most of 
the length of the rostrum with a narrowing gap between them, fusion presumably 
occurring in the premaxillary region at the tip of the rostrum (now lost) . They are 
slightly hollowed ventrally and the external edge projects ventrally to form the 
tomium of the upper mandible. The palatines arise near the posterior ends of the 
maxillae and extend back as shafts of similar width to the maxillae, but tapering 
a little, and then expanding in the region below the orbits, to form elongated, 
ventrally-hollowed blades, their posterior ends articulating with the pterygoids. 
The posterior external sides of both blades are a little broken but together they show 
the line of the external and posterior edge. 


The palato-maxillae arise on the internal edge of the palatines near the region 
where the latter diverge from the maxillae. They extend posteriorly as narrow 
blades lying alongside the internal edge of the palatines, but from the side view of 
the skull it can be seen that they are laterally flattened blades with the upper edges 
curving dorsally and externally on either side of the vomer. Laterally, in the angle 
between the nasal bars and the posterior end of the maxilla, this upper edge of the 
palato-maxilla can be seen on the right side of the specimen as a projecting flange 
level with the junction of the nasal and maxilla ; on the left side it is broken, re- 
vealing the narrow extensions where it joins the internal ventral and internal dorsal 
ends of the maxilla. 

Dorsally, the internal edge of the palatines extends upwards on either side of the 
anterior ventral edge of the interorbital septum, its posterior edge curving inwards 
and downwards to meet the posterior dorsal surface of the vomer. This structure 
is fairly complete but a little broken along its upper edge on the left side, and more 
extensively broken, but still showing curvature, on the right side. 

The pterygoids are slender shafts. In ventral and lateral view the anterior end 
which articulates with the posterior tip of the palatine can be seen to be expanded, 
the sides diverging fairly evenly towards the tip on the ventral surface and with a 
prominent rounded dorso-external end visible laterally. The main shaft shows some 
lateral flattening and where it articulates with the right quadrate it shows dorso- 
external torsion and a rounded, blade-like expansion at the posterior dorsal end. 
This appears to increase the extent of the articulation with the internal side of the 
ventral end of the quadrate. 

Rostrum. Where it joins the skull the rostrum is almost square in transverse 
section, but slightly narrower dorsally ; anteriorly it tapers very gradually to a point. 
The frontal region of the skull terminates in a bold ridge, bordered anteriorly by the 
deep, transverse fronto-nasal hinge. From the base of this ridge a rather flat nasal 
surface of the basal rostrum projects anteriorly, merging into the premaxilla, and 
tapering and becoming more rounded dorsally towards the anterior end. On either 
side it is separated from the more ventrally situated maxillae by an elongated 
aperture which also tapers and which extends to near the distal tip of the rostrum. 
Two slender bars of the nasal slope posteriorly upwards from the posterior part of 
the maxillae just where these widen, and appear to fuse laterally with the edges of 
the broad posterior rostral surface just anterior to the fronto-nasal groove. 

Mandible (Fig. 6f ; PI. 2, figs A-C). The mandibular rami are slender, elon- 
gated and with only a gradual taper towards the tip. They have been twisted and 
show a dorsal torsion to the right. The distal end is damaged but the more proximal 
and articular regions are complete. The dentary portion of each ramus is moder- 
ately rounded externally, more marked towards the ventral edge. The internal side 
of the dorsal edge slants ventro-laterally, and ventral to this a deep groove occupies 
the centre of the internal surface, extending along the distal two-fifths of the man- 
dible. Proximal to this groove the internal surface is rounded with a more promi- 
nent central ridge, continuing the internal tomium edge and gradually descending 



Fig. 5. Views of right coracoid, x 2. A, dorsal ; B, external ; C, internal ; D, ventral. 
Abbreviations : gf, glenoid facet ; pre, procoracoid ; sci, sternocoracoidal impression ; 
sf, sternal facet. 


ventrally and becoming shallower, to disappear near the ventro-posteriorly slanting 
mandibular suture. On the internal side (Fig. 6f), the upper edge of the splenial 
extends about three-quarters of the way along the ventral anterior edge of the 
suture and then extends anteriorly and ventrally as a thin line. 

The mandibular suture appears as a deep groove internally, and much of the 
lower half is occupied by a narrow, elongated fossa, at the anterior end of which a 
narrow slit forms the internal opening of a poorly-defined anterior mandibular 
fenestra. Externally (PL 2, fig. C) the upper part of the suture is fused, but there is 
a narrow, elongated fossa, the dentary edge ventral to it projecting slightly so that 
the fossa is internal to it rather than dorsal. The upper edge of the fossa forms an 
external opening for the fenestral slit. Posterior to this external fossa the suture 
extends back ventro-posteriorly between the dentary and the anterior ventral parts 
of the supra-angular and angular. From the anterior end of the fossa a shallow and 
broad but well-defined groove extends almost horizontally along the external face 
of the dentary and terminates at a level a little posterior to that of the posterior end 
of the groove on the internal face of the mandible. 

On the broad, internal face of the posterior part of the mandible, towards the 
dorsal edge, there is a large, oval, posterior mandibular fossa between the pre- 
articular and supra-angular. Since this is filled with matrix the internal structure 
is not visible, but it does not penetrate to the external surface. 

Posterior to this fossa the mandible becomes less deep, but broadens rapidly just 
before the surface of the articulation with the quadrate. Dorsally, the surface 
widens rapidly into a triangular, smooth surface with a small articular prominence 
at the external corner. On the posterior side of the surface is a deep hollow which 
articulates with the ventral head of the quadrate. It has a complete dorsal rim on 
the external side, but elsewhere slopes to a median groove which occupies the centre 
of an extension of the hollow opening into the internal side. Posterior to the hollow 
the dorsal surface terminates as a narrow edge, broadening at the internal end into 
a small triangular surface with a raised and rounded internal tip, and on the inner 
side of this, broadening the posterior edge of the hollow, there is a small rounded 

The posterior surface of the mandible is flattened and forms a modified inverted 
triangle. It has a distinct and narrow dorso-internal protrusion, while the lowest 
part is more broadly rounded and projects as a narrow, curved lip beyond the main 
ventral shaft. The surface is hollowed at the centre. The true position of this 
surface is difficult to determine because of the torsion of the specimen, but it appears 
to show some internal deflection and a marked posterior-dorsal tilt, the ventral lip 
projecting beyond the rest of the structure. 

b. Postcranial elements 

Sternum (Fig. 6a-c). The sternum was previously concealed within the matrix 
and its presence was not suspected until the present preparation had begun. It is 
still partially embedded in matrix and lacks its posterior end and the lateral pos- 
terior parts of the sternal plate. The carina, which lacks the posterior end, is 
exposed on its right side and along the anterior and ventral edges. In addition the 




Fig. 6. Views of partly embedded sternum, x i ; a. right lateral, b. anterior, c. ventral- 
Views of imperfect left tibiotarsus, x i ; d. oblique anterior, e. oblique internal, f. View 
of the internal side of right mandible, x i . Abbreviations : art, articular ; c, carina ; 
ca, carinal apex ; cs, coracoidal sulcus ; dg, dental groove ; dms, dorsal manubrial 
spine ; irs, inter-ramal suture ; mf, mandibular foramen ; occ, outer cnemial crest ; 
pra, prearticular ; sur, surangular ; vms, ventral manubrial spine. 


ventral manubrial spine and right half of the manubrium and coracoid sulcus 
are also present. The right half of the sternal plate is shattered posterior to the 
thickened region of the sulcus and lacks the external and posterior parts. 

The sternal plate shows slight ventral curvature. The carina is large, projecting 
ventrally and anteriorly, with a slight curvature of the ventral margin. It becomes 
thicker anteriorly, and at the carinal apex bifurcates to accommodate the ventral 
end of the anterior articulating surface. The anterior carinal margin is stout, but 
tapers at the dorsal end where it curves up to the underside of the ventral manubrial 
spine. The latter is a small sharp projection, an inverted triangle in transverse 
section with the wider dorsal surface rising slightly anteriorly and then curving 
forwards and downwards, tapering away to a point a little below the level of the 
ventral edge of the spine. 

The anterior carinal margin curves forwards ventrally, its apex anterior to the 
tip of the spine. The upper part, about three-fifths of the whole, curves forwards 
and becomes thicker ventrally, and forms a blunt projection at a similar anterior 
level to the tip of the manubrial spine. Below this, the remaining two-fifths form 
a flattened, elongated anterior facet, hollowed centrally and curving forward ven- 
trally. It appears analogous to similar surfaces on the sterna of some pelecaniform 
species, which articulate with the furcula in the region of the ventral symphysis. 
Viewed laterally (Fig. 6a), this surface on Prophaethon is hollowed to such an extent 
that it shows some posterior curvature. Viewed anteriorly (Fig. 6b) it is dorso- 
ventrally elongated and wedge-shaped, widening ventrally and with the ventral 
portion curving anteriorly. At the ventral end the hollow becomes shallower, 
forming a poorly-defined lip below the deepest part of the cavity. 

The dorsal lip of the coracoid sulcus appears thickened but is superficially damaged. 
The ventral lip arises at the lateral dorsal edge of the base of the ventral manubrial 
spine and slants posteriorly across the sternal plate. There is a broad ventral 
surface between the anterior edges of the dorsal and ventral lips. The ventral 
labial prominence occurs about two-thirds of the way along the sulcus as a bluntly 
rounded, thin flange overlapping the sulcus which up to this point appears to be 
ventrally exposed. A low ridge across the sternal plate from the posterior part of 
the carina terminates at the sulcus after crossing the ventral surface of the ventral 
labial prominence. 

The sulcus terminates a little lateral to the prominence, the ventrally projecting 
ridge of the dorsal lip curving posteriorly towards the end of the sulcus to leave a 
thinner, laterally-projecting, area of the sterno-coracoid impression. 

Coracoid (Fig. 5 ; PI. 2, figs D-G). The right coracoid is preserved, but an- 
teriorly it is broken off just above the glenoid facet, the broken surface extending 
along part of the ventral edge. The sterno-coracoid process and internal distal 
angle are also damaged. 

The shaft is smooth and rounded, but towards the distal (sternal) end it is dorso- 
ventrally flattened. The sterno-coracoid surface is large and slightly curved 
ventrally, and since the dorsal lip of the coracoid sulcus is markedly anterior to the 
ventral Up, as already described in discussing the sternum, the ridge marking the 


articulating surface for the latter appears across the middle of the dorsal surface of 
the sterno-coracoid area. It becomes lower and disappears before reaching the 
flattened and rather rectangular sterno-coracoid process. The internal distal angle 
of the coracoid is broken off at the line of the articulation ridge. 

The shaft becomes thicker and more rounded, and laterally narrower, as it ap- 
proaches the procoracoid. From the faint scar of attachment of the coraco-brachialis 
a poorly-defined ridge crosses the shaft to the distal end of the procoracoid. 
The latter projects laterally and curves anteriorly and ventrally to a point (broken 
short in the specimen). On the internal side there is a curved hollow between the 
procoracoid and shaft, with a small longitudinal ridge on either side of it. The 
internal opening of the coracoid fenestra is a small hole near the distal end of this 
hollow, and the external opening is a similar hole on the dorsal surface of the pro- 
coracoid, towards its edge. 

The scapular facet is shallow and dorsal, and from it a thickened ridge borders the 
proximal edge of the procoracoid. The glenoid facet is dorsoventrally aligned. 
Ventrally, there is a low ridge from the ventro-exterrial edge of the shaft, which 
crosses the sterno-coracoid process and terminates near its distal external extremity. 

Scapula. The single left scapula lacks the proximal articulating end. It is a 
slender rod with a slight ventral curvature, dorso-v^ntrally flattened. It is a little 
thicker and more rounded at its base and towards its tip becomes more flattened and 
broader, then tapers to a point. 

Pelvis (PI. 3, figs A-C). Most of the pelvis is present. At the anterior end of it 
a row of laterally crushed vertebrae form the relic of the thoracic part of the vertebral 
column. Dorsally the pelvis has been cleared of matrix but some of the more 
ventral detail of the left side is still obscured by matrix containing broken ribs. 
On the right side it is completely exposed. The anterior lateral edge of the ilium 
is broken on the right, but appears complete on the left side. Posteriorly, on the 
right, the ischium and pubis are broken off at about the posterior end of the ilio- 
ischiatic fenestra, and the ilium posterior to the fenestra is also missing, the dorsal 
end terminating at the incomplete caudal end of the synsacrum. On the left side, 
part of the iliac portion of the synsacrum is visible and the ventral struts of the roof 
of the renal depression can also be seen on the left. 

At the anterior end of the pelvis the median-dorsal ridge is thick and prominent, 
appearing to fuse with the end of a row of elongated and flattened neural spines. 
The ridge projects sharply, the iliac plates sloping down steeply on either side and 
curving outwards to form a lateral flange, broadest anteriorly and with a rounded 
tip. Posteriorly the dorsal ridge widens and the iliac plates become narrower and 
less hollowed laterally. The edges of the ridge diverge as two well-defined edges 
of the anterior iliac crest, curving outwards to a slight prominence above and internal 
to the acetabulum. From there, two blunt, broad ridges continue posteriorly as the 
posterior iliac crest, curving slightly towards each other in the region of the ilio- 
ischiatic fenestra, before diverging posteriorly. The shield area between them is 
narrow and slightly hollowed, with a median ridge beginning to appear towards the 


posterior end ; on the left side at the posterior end there is what may have been 
the first of a double row of narrow fenestrae from the renal depression. Below the 
posterior iliac crests the iliac surface slopes outwards as a narrow upper edge to the 
iliac-ischiatic fenestra. The fenestra is large and elongated, becoming wider 
posteriorly, with a relatively straight lower edge, a posteriorly-curved upper edge, 
and the two meeting anteriorly in a small rounded end just posterior to the lower 
edge of the antitrochanter. Anteriorly the surface bordering the upper edge of the 
fenestra flares out to a prominent lip over an antitrochanter, the articulating surface 
of which has an anterior/external aspect and also an anterior/ ventral slant. Below 
this is a rounded acetabulum, the lower third of which does not penetrate completely 
to the ventral side, but forms a rounded hollow in the bone. There is a small 
lateral projection midway along the anterior external edge of the acetabulum. 

Just ventral to the anterior edge of the acetabulum a small anteriorly-projecting 
pectinal process is present, posterior to which the pubis slopes away as a slender rod 
of bone. The anterior end of the ischio-pubic fenestra is narrow but rounded and 
terminates posterior to the ventral edge of the acetabulum. Dorsal to it the ischial 
bar is nearly twice the width of the pubis. Just posterior to the acetabulum the 
external surface of this ischial bar slopes ventro-internally, until it meets a small 
ridge which slopes back from the posterior edge of the antitrochanter, at which 
point torsion occurs and the external face of the ischium has a ventro-external slant, 
and shows a poorly-defined, posteriorly and yentrally slanting, ridge in its surface. 

The fused synsacrum has its greatest depth at the anterior end and viewed laterally 
the dorsal line of the pelvis and the ventral surface of the synsacrum converge 
towards the posterior end, forming a thin elongated wedge that shows a slight ventral 
curvature at its posterior tip. The ventral surface of the synsacrum is widest at the 
region of the sacral vertebrae, where it shows a well-defined median groove, and 
tapers gradually towards the posterior end. Anteriorly it also begins to taper but 
becomes broader again at the anterior tip where it articulates with the first free 
vertebra. The latter appears to be the last dorsal vertebra. The anterior articu- 
lation surface of the synsacrum is posterior to the broad dorsal anterior edge of the 

Femur. The right femur was exposed in the specimen as originally prepared ; 
it appears from the original figure (Andrews 1899) to have had a coating of matrix 
and to have been superficially damaged beneath this. The present preparation has 
exposed on the distal half, which is detached from the main block but still attached 
to the tibiotarsus, an internal distal surface almost to the condyle ; on the proximal 
half still in situ on the pelvis the external and posterior surfaces are visible. 

The distal internal face shows a smooth, slightly rounded surface with a rounded 
and posteriorly projecting condyle, the internal condylar surface showing an internal 
deflection towards the distal end. 

The trochanter shows an abrupt obturator ridge with a hollow beneath it on the 
posterior side. The anterior head of the trochanter is rounded, and the external 
edge appears to form a blunt, projecting ridge which extends for some way along the 
anterior external edge of the shaft. 


TiBiOTARSUS (Fig. 6d-e ; PL 3, figs D-E). During preparation the proximal end 
of the left tibiotarsus was removed and cleaned. The external side of the shaft had 
originally broken away as far as the head, but the internal side is still intact. 

The anterior and internal sides of the shaft are present and where they join there 
is an abrupt edge which becomes more prominent towards the proximal end and 
finally forms the anteriorly-projecting flange of the inner cnemial crest. The inner 
cnemial crest is a thin prominent flange arising along the internal edge, projecting 
anteriorly and at its outer edge curving a little externally. It extends proximally 
well beyond the articulating surfaces. It arises gradually from the shaft, is deepest 
at about the level of the proximal edge of the articulating surfaces, and then tapers 
to a blunt point proximally. The outer cnemial crest arises on the anterior surface 
near the external edge and nearer the proximal end than does the inner crest. It is 
thicker distally than the inner cnemial crest and projects at an angle between the 
anterior and external surfaces, curving slightly towards the external side. Its 
proximal end terminates at an angle a little proximal to the articulation surfaces. 
It appears to end abruptly as though broken short but this is also apparent on some 
entire examples of Recent species. 

The internal articular surface is more distally placed than the inter-articular area 
and the ends of the crests. Its surface is damaged on the specimen, but it shows a 
posteriorly-projecting, curved lip. The inter-articular area slopes distally on the 
external side, curving distally to the broken external surface in a shallow hollow on 
the external side of the outer cnemial crest. 

Ribs and other elements. The head of a rib and part of another are visible on 
the left side of the specimen anterior to the pelvis, together with broken shafts still 
embedded in matrix. The visible head is relatively stout ventral to the tubercle, 
and the portions of shaft are fairly broad. There is no obvious uncinate process, 
but a slender and flat strip of partially embedded bone parallel to the left posterior 
iliac crest might be referable to this. In general structure the ribs resemble those 
of the larger larids and sulids. 

In addition to the material described above, there are also a number of fragments 
of bone and matrix removed during preparation. A close examination of these 
might make their identification possible, but they are not likely to provide additional 
useful information to the present study. Their existence is therefore noted, but no 
further study has been made. 

c. Measurements 

All dimensions are given in millimetres. 

Overall length (premaxilla-supraoc- Maximum width at postorbital pro- 

cipital) 112 cesses 38 

Maximum width at posterior end Minimum width of interorbital bar 15 

(squamosals) 35-5 Width at anterior end of f rentals 16-7 


Skull (continued) 

Width at base of rostrum (dorsal) 

Width at base of rostrum (ventral) 

Mid-rostral width 

Width at tip of rostrum as preserved 

Length of culmen 

Length of lateral nasal aperture 

Maximum depth of lateral nasal 

Posterior depth of cranium to occipital 

Maximum cranial depth 

Depth from anterior end of frontals to 

Width of temporal fossa at postorbital 

Minimum width of cranium at tem- 
poral fossae 


Length of dorsal hollow in orbit 


(? nasal gland) 



Width of dorsal hollow in orbit 





Length of palatines 



Maximum proximal width of palatines 


Length of pterygoid 



Width of shaft of pterygoid 

Width at quadrato-pterygoid articu- 






Maximum width of posterior end of 

basiparasphenoid plate 



Width of rostroparasphenoid 


Length of zygoma 



Depth of zygoma 


Width of zygoma 




Quadratojugal socket to otic process 
External -internal width of quadrato- 

articular surface 
Anterior -posterior width of quadrato- 

articular surface 



External -internal thickness of shaft 

below orbital process 4-8 

Maximum depth of orbital process 6-2 

Length of orbital process 7 • 8 

Lower Mandible 

Maximum length of left dentary to 

post-articular process (incomplete) 109-9 
Maximum length of right dentary to 

post articular process (incomplete) 112-2 
Depth of dentary at tip 4 

Width of dentary at tip i -9 

Depth of dentary at proximal end of 

internal dentary groove 6-7 

Width of dentary at proximal end of 

internal dentary groove 3-7 

Maximum depth of dentary at intra- 

ramal suture 13-2 

Maximum width of dentary at intra- 

ramal suture 



Overall length (incomplete) 


Maximum distal width 



Overall length 


Maximum distal width 


Width of glenoid facet 


Width of scapula facet 


Width of sternal facet 


Depth at anterior end of articular 

surface 6-5 

Width at anterior end of articular 

surface 9-1 

Depth at posterior end of articular 

surface 9-3 

Width at posterior end of articular 

surface 11 -2 

Maximum length of posterior mandi- 
bular fossa 14-3 

Maximum depth of posterior mandi- 
bular fossa 6-7 

Proximal width 4-9 

Thickness at proximal end 2 • 7 

Width of shaft at coracoidal fenestra 6-8 

Internal -external length at sternal 

facet (left) 20-2 

Proximodistal width of sternocora- 

coidal process (left) 8-5 




Maximum length to ventral manubrial 

spine 51-5 

Length of carinal edge (incomplete) 59 

Carinal apex to dorsal edge of ventral 

manubrial spine 26-5 

Tip of ventral manubrial spine to 

anterior carinal margin 6*3 


Maximum medial length as preserved 67 
Anterior border of ilium to anterior 

edge of acetabulum 35 
Maximum width across antitrochan- 

ters 27-4 
Anterior width from external edge of 

ilium to medial ridge 1 7 

Length of ilio-ischiatic fenestra 23-3 
Maximum depth of ilio-ischiatic 

fenestra 10 

Length of furcular facet 10-9 

Maximum width of furcular facet 6-8 

Tip of ventral manubrial spine to 

ventral labial prominence 22-1 

Width of coracoidal sulcus 5-7 

Minimum distance between antitro- 
chanter and pectineal process 14 

Ventral length of synsacrum 69-5 

Anterior depth of pelvis from median 
dorsal ridge to synsacral thoracic 
vertebra 19-5 

Width of articular facet of first syn- 
sacral thoracic vertebra 7 • 5 


Proximodistal length 


Anterior/posterior thickness at mid- 




Length as preserved 


Articular surface to tip of internal 

cnemial crest 


Maximum width of internal cnemial 



Maximum proximal width at trochan- 
teric ridge 9-4 

Width from internal edge to external 

edge of internal cnemial crest 15-5 

Width to internal edge of internal 

cnemial crest 8-6 

Width to external edge of outer 
cnemial crest 14 


In attempting to place Prophaethon within the framework of avian taxonomy, 
using the characters revealed by further preparation, it has been necessary to 
compare it with osteological material from various Recent taxa. Since the earlier 
claims of pelecaniform similarities were based on skull characters it seemed preferable 
for comparative purposes to begin at the other extremity. 

Proximal end of Tibiotarsus. The obvious characters on this element are the 
prominent inner cnemial crest, which has its widest part just above the level of the 
articulation surfaces and tapers proximally, and the inner cnemial crest forming a 
blunt projection at a similar level. The proximal articular surface slopes posteriorly 
and has a projecting lip with a concave proximal surface at the posterior edge. The 
external edge of the proximal surface curves smoothly over onto the shaft in a 
broad zone where it borders the inner cnemial crest. 

Stercorarius provides a very close match in all aspects, while other Charadriiformes 
also show resemblance to varying degrees. In the Procellariiformes Diomedea 


shows some similarities but lacks the projecting posterior lip, and has a larger and 
more proximally situated inner cnemial crest. There is no obvious resemblance to 
the Pelecaniformes, where Phalacrocorax retains only a short curved outer cnemial 
crest while Sula has small blunt projections. The features are almost entirely 
lacking in other pelecaniform genera. 

Femur. This is long and narrow, the trochanteric crest present as a prominent 
narrow ridge rounded off at the proximal end with a curved muscle scar incised into 
the external surface. In the Charadriiformes the ridge is much more developed, 
both anteriorly and proximally. Macronectes in the Procellariiformes shows 
similarities to the specimen. Sula has a stout, curved femur, the proximal end of 
which shows some similarity to that of Prophaethon, but the trochanteric ridge is low 
and blunt and the head is more ventrally deflected. 

Pelvis. The narrow, elongated pelvis is typical of that found today in birds 
which swim and dive to catch their prey. It bears no resemblance to that of 
Phaethon, which is broad and short. 

The anterior shield of the ilium extends forward only as far as the proximal end 
ol the second synsacral thoracic vertebra, and the third vertebra is unfused. This 
condition is typical of the Charadriiformes but not of the other orders examined 
here. Sula does, however, show a short anterior iliac shield. 

The median dorsal ridge is slightly convex, but posteriorly it is depressed between 
well-defined posterior iliac crests. In this respect the specimen resembles Phoehetria, 
Diomedea and Pufinus among the Procellariiformes. The Alcidae show a similar pro- 
file. In the Pelecaniformes the posterior iliac crests are poorly developed and in the 
more aquatic forms such as Sula and Phalacrocorax the posterior median ridge is 
level with or dorsal to the lateral crests. 

The Pelecaniformes also show more laterally projecting and anteriorly directed 
antitrochanteric surfaces than does Prophaethon, and in the latter the iliac surface 
immediately anterior to the acetabulum is concave and the pectineal process pro- 
minent. The specimen is more similar in these respects to both Procellariiformes 
and Charadriiformes, although among the last the Alcidae show the pectineal 
process greatly reduced or absent. From fine sutures apparent on the specimen the 
ilium would appear not to have been fused with the synsacrum, in this respect 
resembling Recent Alcidae and Procellariiformes rather than Pelecaniformes. 

Vertebrae. The free thoracic vertebrae appear to lack hypopophyses. They 
have rounded concavities laterally, posterior to the prominent anterior costal facet. 
The diapophyses are short, dorsally broad and have a thin middle ridge ventrally 
with a deep concavity at the internal corner of the anterior side. There is a thin, 
tapering anterior process at the distal end of the diapophysis. These vertebrae 
resembles those of the larger Larus species in the Charadriiformes. Those of the 
Procellariiformes are more elaborate with larger concavities and various fenestrae, 
and with hypopophyses ; those of the Pelecaniformes show still fewer similarities 
to those of Prophaethon. 


Sternum. Allowing for the incompleteness of the specimen the carina of the 
sternum is deep and long, extending further back than is the case on Recent Pelecani- 
formes. The ventral lip of the coracoid sulcus is posteriorly situated so that the 
sulcus is ventrally exposed, particularly at the middle region, but towards the outer 
end it undercuts a small but broad labial prominence. 

In the Pelecaniformes the ventral lip of the sulcus extends almost as far anteriorly 
as does the dorsal lip, and the sulcus is a deep, anteriorly-directed groove. In 
Prophaethon the groove is more typical of that found in Charadriiformes and Pro- 
cellariiformes. From the ventral labial prominence a distinct intermuscular line 
slants postero-internally towards the middle of the carina. A similar line is present 
in Charadriiformes, but in the Procellariiformes one line arises on the sulcus internal 
to the labial prominence and crosses the sternum more anteriorly, while another 
rises towards the external end and slants towards the posterior end of the carina. 
In Pelecaniformes there is a faint ridge from the labial prominence apparent in 
Phalacrocoracidae and Phaethontidae. 

The hollow facet for furcular articulation at the anterior tip of the carina is a 
distinctive character in Prophaethon. Structures of this kind are found in some 
Recent species of Pelecaniformes and Procellariiformes. A small articulation 
surface is present on the larger Podicipitiformes. This furcular surface is more 
extensively developed in the Pelecaniformes where the Pelecanidae and Fregatidae 
have the furculum fused to the sternum while Phaethontidae, Sulidae and Phala- 
crocoracidae have the surface developed to varying degrees. Of the Procellarii- 
formes the Pelecanoididae have a sternum with a large furcular articulation facet, 
its ventral edge curved anteriorly. The bifurcation of the carina tip, apparent in 
Prophaethon, is characteristic of many Procellariiformes but not of the Pelecani- 
formes. None of the Charadriiformes show furcular articulation facets on the 

The shape of the manubrial spine in Prophaethon resembles that of Phaethon, but 
this structure differs so markedly in different families and genera that we do not 
regard it as taxonomically useful. 

Coracoid. The shape of the sternal (distal) end of the coracoid is correlated 
with that of the coracoid sulcus. The coracoids of Pelecaniformes show sternal 
facets on both sides at the distal end, articulating with the deep sulcus. Prophaethon 
resembles charadriiform and pelecaniform birds in having a prominent facet across 
most of the dorsal surface ; a small one on the ventral surface towards the external 
end is correlated with the position of the ventral labial prominence of the sternum. 
From the fit of the specimen there is no reason to suppose that the missing internal 
distal angle of the coracoid of Prophaethon would have projected across the midline 
of the sternum, as in Phaethon. 

A stout and dorsally curved internal distal angle to the coracoid is characteristic 
of most Procellariiformes, although less evident in the Pelecanoididae, and in the 
Laridae and Alcidae of the Charadriiformes. Although this part of the specimen of 
Prophaethon is damaged, the general shape of the surrounding bone and the sternal 
fragment of the other coracoid, which is also present, indicates an absence of such 


curvature. The specimen most closely resembles the flatter coracoids of Ibido- 
rhynchus and Haematopus of the Charadriiformes in this respect ; it also resembles 
them in the area of irregular surface on the dorsal side distal to the line of attachment 
of the coraco-brachialis muscle, the proportions of the shaft, the procoracoid, and 
the position of the coracoidal fenestra. 

Lower Mandible. This, although relatively complete, does not give an indica- 
tion of afhnity with any particular taxon. 

The slender ramus, increasing in depth posteriorly and tapering a little at the 
anterior end of the articular portion, has a general resemblance to those of Phala- 
crocoracidae and some Procellariidae. The tapering groove on the external surface 
anterior to the intraramal suture is similar to that of Procellariiformes and Char- 
adriiformes, but a groove of this type is also present in Phaethon. 

The arrangement of the component bones around the external fossa of the mandi- 
bular suture and the shape and position of the fossa itself are most closely paralleled 
by the structure in some smaller Laridae, the Burhinidae and to a lesser degree by 
some Procellariiformes. The concave, ventrally tapering and postero-dorsally 
oriented postarticular surface is very similar to that of Gaviiformes, but only 
resembles to a limited extent those of the other taxa examined here. 

The articular facets are very similar to those of Laridae, Burhinidae and Pro- 
cellariidae but do not closely resemble the more specialized structures of the Pele- 

Prophaethon has a large, rounded posterior mandibular fossa, bordered anteriorly 
by a large prearticular which extends anteriorly to border the mandibular suture 
and overlaps with, and possibly fuses with, the dentary towards the dorsal edge of 
the ramus. This large prearticular is also present in the Procellariiformes and 
Gaviiformes. The Charadriiformes have a large posterior fossa but the prearticular 
is small and only partially occupies the anterior space, the mandibular slit forming a 
distinct fenestra. Phaethon shows a similar condition but has the prearticular 
larger posteriorly, reducing the size of the posterior fossa. In the other pelecaniform 
families the posterior fossa is tiny and the prearticular has filled the remaining 

Skull. The palate is typically schizognathous and resembles those of Charadrii- 
formes and Gaviiformes. Procellariiformes also have this type of palate, but the 
distal end is modified by bill shape. The palates of Pelecaniformes appear to be 
schizognathous in the very juvenile condition but desmognathous in the adults. 
Schizognathous palates are also present in Podicipitiformes, Gruiformes, Galli- 
formes, Sphenisciformes and Columbiformes. The distinctive proximal bifurcation 
of the vomer, visible between the palatines in Prophaethon, is very similar to that of 
larger Larus species and of other Charadriiformes, but does not appear in Pelecani- 
formes and Procellariiformes. 

The rostrum of Prophaethon is long, slender and tapering. An elongated nasal 
aperture runs for almost the whole length, rising posteriorly and tapering to a slit 
just proximal to the frontonasal hinge. In general shape the rostrum is similar 


to that of the Phalacrocoracidae, and if it were argued that the latter retained the 
open aperture of the juvenile there would be strong similarity. In other Recent 
birds the elongated nasal apertures are typical of Charadriiformes, Gaviiformes, 
Podicipitiformes and Gruiformes. Although the nares of Pelecaniformes are 
typically closed or minute, Phaethon shows an intermediate condition with a short 
but relatively large aperture about a third of the way along the rostrum and a tiny 
hole near the frontonasal hinge. 

A transverse frontonasal hinge developed to differing degrees in different families 
is associated with the greatly reduced nostrils in Pelecaniformes. In Prophaethon 
the nasal apertures are schizorhinal in shape, but since the nasal struts join the 
rostrum anterior to the frontonasal hinge they are functionally holorhinal. A 
holorhinal type of nostril associated with a frontonasal hinge in this position also 
occurs in the Charadriiformes in the Burhinidae, Thinicoridae and Pluvianus of the 
Glareolidae, although charadriiform nostrils are usually collectively described as 
schizorhinal. A deep transverse frontonasal hinge comparable with that of the 
Phaethontidae also occurs in Rhynchops of the Charadriiformes, but in this instance 
the nostrils are schizorhinal. Procellariiformes have an unspecialized transverse 
frontonasal structure and holorhinal nostrils. 

In the Pelecaniformes the heavy frontal brow associated with a deep frontonasal 
groove is present only in the Phaethontidae. In the Charadriiformes it is partially 
developed in Rhynchops, and more highly developed in the Chionidae. In the 
latter the nasal struts lie alongside the rostrum and also terminate just below this 
brow, apparently forming a hinge with the rostrum. This structure has some 
analogy with that of Prophaethon since a close inspection of the latter reveals that 
the nasal lies close alongside the dorsal rostral surface but may not fuse with it 
completely, and the transverse line of the frontal brow is not completely straight but 
shows paired lateral recesses in the fore-edge which might be comparable with those 
of Chionidae. Unfortunately the surfaces of the bone are slightly damaged at this 
point in the specimen oi Prophaethon. 

The frontal shows a flattened lateral surface suggesting the loss of an unfused 
prefrontal. Prefrontals are unfused in the Phaethontidae and Fregatidae of the 
Pelecaniformes, and in Burhinidae of the Charadriiformes ; when they become 
detached they leave fiat surfaces similar to those of Prophaethon. 

The smooth dorsal surface of the skull of Prophaethon may have pelecaniform 
similarities, but only by virtue of the absence of the nasal glands, and were these not 
present in other taxa they might also be similar. The feature is therefore a general- 
ized one, indicative of absence of specialization rather than affinity. In the deeply 
rounded temporal fossae posterior to the prominently curved postorbital process 
the specimen resembles the Fregatidae, Uria of the Alcidae and Stercorarius in the 
Laridae, rather than the Phaethontidae, but shows some similarity to the last in 
the rather limited development of the fossae dorsally and anteriorly. 

In the area of quadrate articulation the ventral edge of the orbitosphenoid curves 
forwards, leaving a large cavity anterior and internal to the articulation surface and 
tending to divide the latter into two. This is also present to a similar degree in the 
Pelecaniformes, but much reduced in the Procellariiformes and Charadriiformes. 


The posterior aspect of the skull of Prophaethon has parallels in all the major taxa 
here examined, and there are considerable differences in these at the family level. 
The supraoccipital of Prophaethon has a small foramen on either side towards the 
ventral edge, with associated grooves bordering the upper edge of the foramen 
magnum. Similar structures are apparent in the Procellariiformes, more modified 
in the Charadriiformes, and greatly modified or absent in the Pelecaniformes. 

Within the orbit of Prophaethon the hollow apparently accommodating the nasal 
gland is in a unique position, being an elongated hollow bordering the dorsal edge of 
the orbit and on the ventral surface of it. In Pelecaniformes the nasal gland is 
usually towards the anterior dorsal end of the orbital hollow and in a more median 
position. In some Phaethon species a small elongated hollow extends posteriorly 
into the orbit from this anterior site, bordering the orbital septum but separated 
from it by the channel of the olfactory nerve. In the Sulidae it is an elongated 
hollow lying completely in the roof of the orbit, but still close to the septum. In 
Charadriiformes and Procellariiformes the nasal gland hollow either borders the 
dorsal edge of the orbit or lies internal to it, but on the dorsal surface of the frontal. 
The position of this structure in Prophaethon is therefore intermediate between that 
in the two Recent types. 

The slender, unspecialized pterygoids of Prophaethon, with a small posterior 
lateral expansion of the orbital process of the quadrate, give little useful evidence of 
affinity. The Phaethon tidae show an even less modified structure, but the ptery- 
goids of other pelecaniform families and of the other taxa discussed here are stouter, 
with some development of lateral flanges on the shaft, and in several diverse taxa 
show the expanded posterior end. 

The quadrate of Prophaethon also shows little evidence of affinities, and the deep 
groove at the ventral end of the posterior surface of the shaft appears to be a pecu- 
liarity of the species. 

Tables i and 2 give a simplified summary of apparent similarities. Tables such 
as this may oversimplify, exaggerating both similarities and differences, but they 
indicate the problem involved in assigning Prophaethon to a known taxon. 

Table i 

Comparison of postcranial elements of Prophaethon with extant orders of birds 

Charadriiformes Procellariiformes Pelecaniformes 

Proximal end, tibiotarsus * x - 

Proximal end, femur — x x 

Pelvis * * X 

Vertebrae * x - 

Sternum, other than anterior facet * x — 

Anterior edge of carina _ * * 

Coracoid * * x 

* very similar; x some similarity; - little or no similarity 


reappraisal of 
Table 2 

Comparison of cranial elements of Prophaethon with extant orders of birds 

Charadrii- Procellarii- Phaethon- 
formes formes tidae 

Lower Mandible 


External intraramal suture 






Posterior fossa 









Frontonasal hinge 
Unfused prefrontals 
Temporal fossae 
Supraoccipital area 
Quadrate articulation area 





* very similar; x 

some similarity; 




little or no similarity 


From the above data comparing Prophaethon with Recent birds there would appear 
to be several possible hypotheses concerning its relationship to known taxa. 

The species might be regarded as a charadriiform bird, showing the alcid type of 
pelvis and with a head adapted for catching prey in water and showing modifications 
which have parallels elsewhere in the taxon. But the specialized sternum with its 
facet for furcular attachment is not present in known Charadriiformes, and it would 
be necessary to argue that, since it is known in aquatic birds of two other orders, it 
might be an adaptation that potentially could have evolved under any similar 
behavioural or environmental selection pressures. 

Prophaethon might be regarded as a procellariiform bird, in which case there is a 
precedent for a modified sternum, but the skull and rostral characters would not be 
typical of the known forms of that order. 

If it is argued that the possession of both the specialized frontonasal hinge and 
the sternum are indications of pelecaniform affinities, then another assumption must 
be made, that a saltatory form of evolution has produced the more extremely 
evolved characters now present in this evolutionary diverse order while retaining 
more generalized characters in other structures which could presumably undergo 
subsequent modification. This is at variance with the apparent evidence provided 
by a proto-frigatebird from the Lower Eocene, described by Olsen (1974), which 
shows modifications in various skeletal elements of the postcranial skeleton but has 
not achieved either the specialized skull or the sternum typical of the Fregatidae. 

If any of the above suggestions are adopted then, at least in so far as the early 
Tertiary is involved, the range of osteological characters used to define any of the 


three orders must be extended to include a number of significant characters which 
are at present regarded as diagnostic of some other order. The Hmiting groups of 
characters which may be used to assign any species to a particular order will then 
no longer be clearly separable ; we shall have to assume it is not possible to identify 
a bird from this geological period unless certain critical skeletal elements are avail- 

Alternatively, in view of the lack of evidence of affinity with any single order, it 
could be suggested that in the early Tertiary these three Recent orders had not yet 
diverged, and that Prophaethon was referable to an ancestral stem from which 
more than one Recent order had subsequently evolved. There is, however, a 
considerable amount of fossil bird material from this period, most of which appears 
to be referable to Recent families, suggesting that the degree of evolutionary diver- 
gence apparent in Recent families had already occurred by this period. In addition 
fossils referred to Pelecaniformes and Charadriiformes are known from the Upper 
Cretaceous and from the evidence available it seems that in general the ordinal 
divisions of birds had occurred within the Cretaceous. 

Some other Lower Eocene species show this intermediacy of affinities. Among the 
more specialized forms described from this period is a sea-bird with bony, tooth-like 
projections on the jaws, Odontopteryx toliapica of the family Odontopterygidae. 
Related forms, usually separated in the family Pseudodontornithidae, are known 
from the Miocene. In recent studies of the British Lower Eocene we have found 
evidence of more numerous and varied forms of both families at this period. These 
birds were at first regarded as pelecaniform, but Howard (1957), in describing 
Osteodontornis orri of the Californian Miocene, pointed out that the skeleton showed 
a mixture of pelecaniform and procellariiform characters. The British specimens, 
recently prepared, appear to confirm the rather specialized nature of this group. 

Avian phylogeny in the Cretaceous is still a mystery and in view of the lack of 
evidence speculation is of very limited use. If the Lower Eocene is taken as a base 
line there is an array of families known from fossil remains, most of them first known 
to occur at this period and subsequently persisting until Recent times. If we 
insert into this array both Prophaethon and the bony-toothed birds we may produce 
the kind of picture shown in simplified and linear form in Fig. 7. 

Let us assume that the base line is the Lower Eocene and the lines rise to the 
Holocene, that A1-A3 represents charadriiform families, B1-B3 pelecaniform 
families and C1-C3 procellariiform families. If we insert Prophaethon as AB, the 
Odontopterygidae as BCi and the Pseudodontornithidae as BC2, then we have a 
continuous sequence in which each family shares some characters with the adjacent 
ones and the whole presents a relatively uniform array which might have arisen as a 
complex adaptive radiation from a single ancestral origin, rather than as a divergence 
from three separate stems, yet showing some morphological convergence. If during 
later periods Prophaethon and the bony-toothed birds became extinct, then the 
remaining families would appear to fall into more discrete groups which are then 
identified as Recent orders - ordinal definition being aided by the absence of the 
intervening forms. The diagram presented is a linear one, but the relationships 
should be visualized as three-dimensional, the three Recent orders forming a triangle. 


Bl B2 B3 



Fig. 7. Hypothetical relationship of families. 

The bony-toothed birds would occupy the space intervening between the Pelecani- 
formes and the Procellariiformes, while Prophaethon would come in a more central 
position in view of its similarities to all three orders. 

We have also indicated by transverse lines two possible distributions of shared 
characters resulting from the loss of some of these families. If BC1-2 disappear 
then the two linking characters which they share become discrete ordinal peculi- 
arities. The character which AB is shown as sharing with families on either side 
would, following the extinction of AB, still persist in more than one Recent order 
but might be limited to certain families. If we assume A1-3 to be charadriiform 
families and Bi to be the Phaethontidae, species of the latter are seen to share a 
number of minor morphological characters with the former group and not with other 
pelecaniform families. The existence of such characters led to some early sug- 
gestions that the Phaethontidae had more affinity with the Charadriiformes than 
with the Pelecaniformes, and Mathews & Iredale (1921) created for it a suborder 
within their gull order, Lari. These similarities have also been explained as con- 
vergence with the terns (Sterninae of the Laridae) brought about by similar methods 
of feeding. The latter might be a valid explanation, but on the basis of the hypo- 
thesis proposed some at least of these similarities might be explained by a closer 
degree of affinity between the families involved at an earlier period and by the 
persistence of some shared characters. 

This proposed arrangement would appear the most satisfactory to explain the 
combinations of characters found in Prophaethon and the bony-toothed birds, and 
to present a balanced view of sea-bird evolution in the Lower Eocene based on 
present evidence. There has for long been general acceptance that the pelecaniform 
and procellariiform birds shared a common ancestry more recent than that of avian 
orders in general. The suggested association of the Charadriiformes with these has 
been weaker and based mainly on the peculiarities shown by the Phaethontidae. 


The evidence from the skeletal structure of Prophaethon might now justify, pro- 
visionally, a stronger linking of the Charadriiformes with the Pelecaniformes. 
These three Recent orders, with the two interordinal linking taxa, appear to form a 
more unified group, perhaps a single superorder which, for the purposes of arranging 
avian taxa of the Lower Eocene and earlier periods, might provide a more convenient 
unit than the existing groups. 

This idea does not significantly conflict with present views on affinities based on 
the study of Recent taxa. These are summarized by Sibley & Ahlquist (1972) in 
their study of non-passerine relationships based on egg-white protein electrophoresis. 
There are various characters which appear to link the three Recent orders, and in 
addition the Gaviiformes are considered to be closely related to the Charadriiformes 
and the Sphenisciformes to the Procellariiformes. These two additional orders 
should perhaps be included in any proposed major grouping. Sibley & Ahlquist 
found evidence of similar starch gel egg-white patterns in Sphenisciformes, Gavii- 
formes, Procellariiformes, Charadriiformes, and possibly the Pelecanidae, Fregatidae 
and Phaethontidae. There is therefore evidence of a large and interrelated supra- 
ordinal group, within which some earlier links may still be undiscovered. 

While this hypothesis may help to establish the position of Prophaethon within the 
framework of palaeontological phylogeny, the problem of fitting it into the existing 
taxonomic hierarchy still remains. As we have already indicated, it does not show 
definite affinity with any single existing taxon and to attempt to associate it with 
one of these would affect ordinal definitions based on osteology. From a nomen- 
clatural point of view the alternative treatment of the specimen seems preferable 
whereby its isolated position is recognized. In terms of Recent taxonomy Pro- 
phaethon constitutes an interordinal link. For nomenclatural purposes it seems 
preferable to treat it as a monotypic order, Prophaethontiformes, with the single 
family Prophaethontidae, while recognizing its special character. It is then possible 
to insert it into the existing framework without unduly affecting existing taxa, and 
it can be merged with another taxon at a later date if new and overriding evidence 
should justify this. For the moment we would recommend that its placement 
should be between the Pelecaniformes and the Charadriiformes. 


Ordinal Diagnosis. Dorsal surface of skull with prominent postorbital process 
and deep, rounded temporal fossae. Cranium small. Anterior lateral surface of 
frontal flattened for attachment of unfused prefrontal. Deep fronto-nasal hinge 
with prominent brow ridge. Lateral nasal struts He alongside, and may fuse with, 
rostrum just anterior to hinge. Elongated, oval depression in roof of orbit bordering 
external edge. Zygoma laterally flattened with no dorso-ventral torsion at anterior 
end. Rostrum tapering evenly with nares approaching schizorhinal condition but 
terminating anterior to fronto-nasal hinge. Elongated nasal aperture extending 
almost to rostral tip. Palate typically schizognathous. Quadrate with large 
projecting flange on otic process, and posterior lower end of shaft with deep, narrow 


groove. Lower mandible long and slender. Dentary with horizontal external 
groove. Posterior mandibular fossa large, not perforating external wall. Pre- 
articular large. 

Carina of sternum deep, with curved ventral edge and anteriorly projecting apex 
with well-developed manubrial spine. Flattened anterior surface at carinal apex 
for articulation of furcula, and carinal tip bifurcated. Coracoid sulcus shallow with 
poorly-developed ventral lip. Ventral labial prominence projecting and rounded. 

Coracoid with large sterno-coracoid surface, slightly curved ventrally, and sternal 
facet wide. Small facet for labial prominence on ventral surface. Procoracoid 
stout, projecting laterally and curving anteriorly. Coracoid fenestra present. 
Glenoid facet projecting dorso-ventrally. 

Pelvis elongated and relatively narrow, with prominent median dorsal ridge, and 
narrow posterior shield between raised posterior iliac crests. Fused synsacrum with 
greatest depth at anterior end, tapering posteriorly. Last thoracic vertebra not 
fused to synsacrum. Anterior iliac plates sloping down steeply and curving outwards 
to form broad, lateral flanges which project a little anterior to end of the synsacrum. 

Femur with internal cotylar surface internally deflected towards distal end. 
Proximal trochanter has abrupt obturator ridge with hollow on posterior side. 
Trochanteric ridge prominent. 

Inner cnemial crest of tibiotarsus extending proximally well beyond anterior 
surfaces, as a thin flange projecting prominently, with some external curvature at 
outer edge. Outer cnemial crest thicker distally than inner crest, with proximal 
end terminating at an angle a little proximal to articulating surfaces. Internal 
articular surface with a posteriorly projecting curved lip. 

Diagnosis. The only family of its order. 

Genus PROPHAET HON Andrews 1899 
Diagnosis. The only genus of its family. 
Type Species. Prophaethon shrubsolei Andrews. 

Prophaethon shrubsolei Andrews 1899 
(Pis 1-3 ; Figs 1-6) 

1899 Prophaethon shrubsolei Andrews : 776-785, pi. 51. 

Diagnosis. The only species of its genus. 

HoLOTYPE. Imperfect skull, lower jaws and hyoids, sternum, right coracoid, 
distal end of left coracoid, left scapula, right femur, proximal end of left tibiotarsus, 
II vertebrae, rib fragments and synsacrum. In British Museum (Natural History), 
Department of Palaeontology registered number A683. 

Locality and Horizon. Lower Eocene, London Clay (Ypresian) of Sheppey, 
Kent, England. 



We wish to thank Mr F. M. P. Howie for preparing the specimen, Miss M. L. 
Holloway for making the detailed line drawings, and Mr F. Greenaway for taking the 


Andrews, C. W. 1899. On the remains of a new bird from the London Clay of Sheppey. 

Proc. zool. Soc. Lond., 1899 : 776-785 ; pi. 51. 
Howard, H. 1929. The avifauna of the Emeryville shellmound. Univ. Calif. Pubis Zool., 

Berkeley, 32 : 301-394. 
1957- A gigantic 'toothed' marine bird from the Miocene of California. Bull. Dep. 

Geol. S. Barbara Mus. nat. Hist., 1 : 1-23, 8 figs. 
JoLLiE, N. T. 1957. The head skeleton of the chicken and remarks on the anatomy of this 

region in other birds. /. Morph., Boston, Mass., 100 : 389-436, 26 figs. 
Mathews, G. M. & Iredale, T. 1921. A Manual of the Birds of Australia, 1. 279 pp. London. 
Olsen, S. L. 1974. New fossil evidence of the origin of Frigatebirds. Abstr. i6th int. orn. 

Congr., Canberra : 61-62. 
Pycraft, W. p. 1898. Contributions to the Osteology of Birds. Part I. Steganopodes. 

Proc. zool. Soc. Lond., 1898 : 82-101, pis 7-8, figs 1-8. 
Sibley, C. G. & Ahlquist, J. E. 1972. A comparative study of the egg white protein of 

non-passerine birds. Bull. Peabody Mus. nat. Hist., New Haven, Conn., 39:1-246, 

37 figs. 


The page numbers of the principal references are printed in bold type. An asterisk (*) 
denotes a figure. 

Alcidae 19-20, 22 
Andrews, C. A. 3-4 

bony-toothed birds 25-6 
Burhinidae 21-2 

Galliformes 21 
Gaviiformes 21 
Glareolidae 22 
Greenaway, F. 29 
Gruiformes 21-2 

2, 27 

canna 23 

Charadriiformes 3, 18-27 

Chionidae 22 

Columbiformes 21 

comparisons with Recent material 18-24 

coracoid 4, 10*, 13-14, 17, 20-1, 23 ; pi. 2, 

figs D-G 
cranial structures 4-8, 24 
Cretaceous, ordinal divisions of birds within 


Diomedea 18-19 

egg-white protein electrophoresis 27 
evolution of birds 24-7 

families, hypothetical relationship of 23, 26* 
femur 4, 15, 18-19, 23 

Fregatidae 20, 22, 24, 27 

Haematopus 21 
Holloway, Miss M. L. 29 
Howie, F. M. P. 4, 29 

Ibidorhynchus 21 

jaw, lower, see mandible 

Lari 26 

Laridae 20-2, 26 

Larus 19, 21 

limb-bones 4 ; see femur, etc. 

Macronectes 19 

mandible, lower 4, 9-11, 12' 

pi. 2, figs A-C 
measurements 16-18 

nasal structures 21-2 
nostrils 4 

17, 21, 24 



Odontopter^^gidae 25 
Odontopteryx toliapica 25 
orbital structure 4-7 
Osteodontornis orri 25 

palate, 4, 7*, 8-9, 21 ; pi. i, fig. B 

Pelecanidae 20, 27 

Pelecaniformes 3, 6, 19-27 

Pelecanoididae 20 

pelvis 4, 14-15, 18-19, 23 ; pi. 3, figs A-C 

Phaethon 4, 19-23 

Phaethontidae 3-4, 20, 22-4, 26-7 

Phalacrocoracidae 20-2 

Phalacrocovax 4, 19 

Phoebetvia 19 

Pluvianus 22 

Podicipitiformes 20-2 

postcranial elements 11-16, 23 

Procellariidae 21 

Procellariiformes 3, 18-27 

Prophaethon shruhsolei 1-29 passim, 28; 

pis 1-3 
Prophaethontidae fam. nov. 27, 28 
Prophaethontiformes ord. nov. 3, 27-8 
proto-frigatebird 24 
Pseudodontornithidae 25 
Puffinus 19 

quadrate 8, 17 

Rhynchops 22 
ribs 4, 16 
rostrum 9, 21-2 

scapula 4, 14, 17 

Sheppey, Isle of 3 

Shrubsole, W. H. 3 

skull 3, 4-11, 5*, 7*, 16-17, 21-4; pi. I, 

figs A-E 
Sphenisciformes 21, 27 
Stercorarius 18, 22 
Sterninae 26 

sternum 4, 11-13, 12*, 18, 20, 23 
Sula 4, 19 
Sulidae 20, 23 

Thinicoridae 22 

tibiotarsus 4, 12*, 16, 18-19, 23; pi. 3, 
figs D-E 

Uria 22 

vertebrae 4, 19, 23 

X-rays 4 

Dr C. J. O. Harrison, Ph.D. 

Subdepartment of Ornithology 
British Museum (Natural History) 

C. A. Walker 

Department of Palaeontology 
British Museum (Natural History) 
, Cromwell Road 
London SW7 5BD 

Accepted for publication 18 July 1975 

Prophaethon shrubsolei Andrews 

BM(NH) Pal. Dept. no. A683. Views of skull, x i. 

A. dorsal (p. 5) 

B. ventral (p. 7, 8) 

C. left lateral (p. 4) 

D. right lateral (p. 4) 

E. posterior (p. 7) 

Bull. Br. Mils. flat. Hist. (Geol.) 27, 


.i.«MMiBuittMMii.^ ':'''^ 

Prophaethon shrubsolei Andrews 

BM(NH) Pal. Dept. no. A683. 

BM(NH) Pal. Dept. no. A683. 

Views of lower mandible, x i. (p. 9) 

A. dorsal 

B. ventral 

C. left lateral 

Views of right coracoid, 

D. dorsal 

E. external 

F. internal 

G. ventral 

(P- 13) 

Bull. Br. Mus. nat. Hist. (Geol.) 27, i 


Prophaethon shrubsolei Andrews 

BM(NH) Pal. Dept. no. A683. 

BM(NH) Pal. Dept. no. A683. 

Views of pelvis, x i. (p. 14) 

A. dorsal 

B. ventral 

C. right lateral 

Views of proximal end of left tibiotarsus, x 2. (p. i( 

D. anterior 

E. internal 

Bull. Br. Mus. nat. Hist. (Geol.) 27, i 







1. Cox, L. R. Jurassic Bivalvia and Gastropoda from Tanganyika and Kenya. 
Pp. 213 ; 30 Plates ; 2 Text-figures. 1965. out of print. 

2. El-Naggar, Z. R. Stratigraphy and Planktonic Foraminifera of the Upper 
Cretaceous — Lower Tertiary Succession in the Esna-Idfu Region, Nile Valley, 
Egypt, U.A.R. Pp. 291 ; 23 Plates ; 18 Text-figures. 1966. £11. 

3. Davey, R. J., DowNiE, C, Sarjeant, W. a. S. & Williams, G. L. Studies on 
Mesozoic and Cainozoic Dinoflagellate Cysts. Pp. 248 ; 28 Plates ; 64 Text- 
figures. 1966. ;f8.20. 

3. Appendix. Davey, R. J., Downie, C, Sarjeant, W. A. S. & Williams, G. L. 
Appendix to Studies on Mesozoic and Cainozoic Dinoflagellate Cysts. Pp. 24. 
1969. 95p. 

4. Elliott, G. F. Permian to Palaeocene Calcareous Algae (Dasycladaceae) of 
the Middle East. Pp. iii ; 24 Plates ; 16 Text-figures. 1968. ;^ 

5. Rhodes, F. H. T., Austin, R. L. & Druce, E. C. British Avonian (Carboni- 
ferous) Conodont faunas, and their value in local and continental correlation. 
Pp- 313 I 31 Plates ; 92 Text-figures. 1969. £13.10. 

6. Childs, a. Upper Jurassic RhynchoneUid Brachiopods from Northwestern 
Europe. Pp. 119 ; 12 Plates ; 40 Text-figures. 1969. 3^5.25. 

7. Goody, P. C. The relationships of certain Upper Cretaceous Teleosts with 
special reference to the Myctophoids. Pp. 255 ; 102 Text-figures. 1969. 


8. Owen, H. G. Middle Albian Stratigraphy in the Anglo-Paris Basin. Pp, 164 ; 
3 Plates ; 52 Text-figures. 1971. ;^7.20. 

9. Siddiqui, Q. a. Early Tertiary Ostracoda of the family Trachyleberididae 
from West Pakistan. Pp. 98 ; 42 Plates ; 7 Text-figures. 1971. £9.60. 

10. FoREY, P. L. a revision of the elopiform fishes, fossil and Recent. Pp, 222; 
92 Text-figures. 1973. £11.35. 

11. Williams, A. Ordovician Brachiopoda from the Shelve District, Shropshire. 
Pp. 163 ; 28 Plates ; 11 Text-figures ; no Tables. 1974. £12.80. 

Printed in Great Britain by John Wright and Sons Ltd. at The Stonebridge Press, Bristol BS4 5NU 

6&n , 


"^ GthtrsAW - 

29 JUL 




GEOLOGY Vol. 27 No. 2 

LONDON: 1976 




British Museum (Natural History), London 


Institute of Systematic and Experimental Zoology, 
Polish Academy of Sciences, Krakow, Poland 

Pp. 31-147 ; 31 Text-figures ; 13 Tables 

GEOLOGY Vol. 27 No. 2 

LONDON: 1976 


(natural history), instituted in 1949, is 
issued in five series corresponding to the Scientific 
Departments of the Museum, and an Historical series. 

Parts will appear at irregular intervals as they 
become ready. Volumes will contain about three or 
four hundred pages, and will not necessarily be 
completed within one calendar year. 

In 1965 a separate supplementary series of longer 
papers was instituted, numbered serially for each 

This paper is Vol. 27, No. 2, of the Geological 
[Palaeontological) series. The abbreviated titles of 
periodicals cited follow those of the World List of 
Scientific Periodicals. 

World List abbreviation : 
Bull. Br. Mus. nat. Hist. (Geol. 

ISSN 0007-1471 

Trustees of the British Museum (Natural History), 1976 


Issued 29 July, 1976 Price £7.40 




Synopsis ........ 

I. Introduction ....... 

A. History of Studies ..... 

B. The Geological Background .... 
II. Localities in the British Isles with fossil rodents 

A. Deposits OF East Anglia . .... 

(i) Red Crag 

(ii) Icenian Crag ....... 

(iii) Cromer Forest Bed Series ..... 

(a) Pastonian of East Runton and Happisburgh . 

(b) Beestonian ...... 

(c) Cromerian, sensu stricto .... 

(d) Anglian ....... 

B. Terrace and Solifluxion deposits of the Rivers Thames 

AND Lea ........ 

(i) Lower Thames ....... 

(a) High Terrace of Swanscombe 

(b) Clacton ....... 

(c) Early Middle Terrace at Aveley, Grays Thurrock 

and Ilford ...... 

(d) Late Middle Terrace at Crayford and Erith . 

(e) Floodplain Terrace complex 

(f ) The Middle Terrace/Floodplain Terrace problem 
(ii) Upper Thames ....... 

(a) Isleworth ....... 

(b) Marlow 

(iii) River Lea ....... 

(a) Water Hall Farm Gravel Pit 

(b) Nazeing, Ponders End, Edmonton and Hackney 
(iv) Solifluxion and melt-water deposits 

(a) Northfleet, Ebbsfieet and Baker's Hole 

C. Cave Deposits ....... 

(i) Westbury Fissure ...... 

(ii) Kent's Cavern ...... 

(iii) Tornewton Cave ...... 

(iv) Other caves with pre-Tpswichian' deposits . 

(v) 'Ipswichian' cave deposits .... 

(vi) Caves with post-Ipswichian deposits . 
(vii) Caves in Scotland ...... 

(viii) Caves in Ireland ...... 

D. Other Localities ....... 

(i) Cromerian localities ...... 














(ii) Hoxnian localities 

(iii) 'Ipswichian' localities 

(iv) Last Glaciation localities 

(v) Holocene localities 

(vi) Scottish localities 

IIL Classification and distribution of rodents in the Pleistocene 
OF the British Isles 
Family Sciuridae Brandt 
Genus Sciurus Linnaeus 

Sciurus whitei Hinton 

Sciurus vulgaris Linnaeus 
Genus Spermophilus Cuvier 

Spermophilus {Urocitellus) primigenius Kormos 

Spermophilus {Coloboiis) super ciliosus (Kaup) 
Family Gliridae Thomas 
Genus Muscardinus Kaup 

Muscardinus avellanarius (Linnaeus) 
Family Castoridae Gray 

Genus Trogontherium Fischer 

Trogontherium minus Newton . 

Trogontherium boisvilletti (Laugel) 
Genus Castor Linnaeus 

Castor fiber Linnaeus 
Family Muridae Gray . 
Genus Apodemus Kaup 

Apodemus sylvaticus (Linnaeus) 

Apodemus flavicollis (Melchior) 
Genus Micromys Dehm 

Micromys minutus (Pallas) 
Genus Mus Linnaeus . 

Mus musculus Linnaeus . 
Genus Rattus Fischer. 

Rattus rattus (Linnaeus) . 

Rattus norvegicus (Berkenhout) 
Family Cricetidae Rochebrune 
Genus Cricetus Leske. 

Cricetus cricetus (Linnaeus) 
Genus Allocricetus Schaub . 

cf. Allocricetus bursae Schaub , 
Genus Dicrostonyx Gloger . 

Dicrostonyx torquatus (Pallas) 
Genus Lemmus Link . 

Lemmus lemmus (Linnaeus) 
Genus Clethrionomys Tilesius 

Clethrionomys glareolus (Schreber) 
Genus Pliomys Mehely 

Pliomys episcopalis Mehely 
Genus Mimomys Major 

Mimomys pliocaenicus Major 

Mimomys reidi Hinton . 

Mimomys newtoni Major. 

Mimomys savini Hinton . 









Genus Arvicola Lacepede ....... 99 

Arvicola cantiana (Hinton) 


Arvicola cantiana-terrestris 


Arvicola terrestris (Linnaeus) 


Genus Pitymys McMurtrie 


Pitymys arvaloides Hinton 


Pitymys gregaloides Hinton 


Genus Microtus Schrank . 


Microtus arvaliniis Hinton 


Microtus agrestis (Linnaeus) 


Microtus arvalis (Pallas) 


Microtus nivaloides Major 


Microtus nivalis (Martins) 


Microtus ratticepoides Hinton 


Microtus oeconomus (Pallas) 


Microtus gregalis (Pallas) 


Genus Lagurus Gloger 


Lagurus lagurus (Pallas) . 


IV. History of the rodent fauna of the British Isles during thi 

Pleistocene ........ 


Red Crag 



Icenian Crag and Pastonian Forest Bed 



Cromerian sensu stricto . 



'Westbury stage' .... 


Mundesley Arctic Bed . 



Hoxnian deposits .... 


Comparison of the rodent faunas . . . 


Notes on the Wolstonian-Ipswichian part of the succession 


Middle Terrace of the Thames interglacial 


Tornewton Cave Glutton Stratum cold stage . 


Joint Mitnor interglacial ...... 


Last Glaciation . 


Holocene .... 


Ireland .... 


V. Acknowledgements 


VI. References .... 


/II. Index ..... 



For nearly half a century, since its publication in 1926, M. A. C. Hinton's Monograph of the 
Voles and Lemmings has remained the only comprehensive work on British Pleistocene rodents. 
Subsequent advances in Quaternary studies and the availability of new fossil material have made 
some updating of this publication necessary. In the present work a brief historical review of 
the studies of British Pleistocene rodents is followed by a description of the rodent assemblages 
from the principal fossil localities. Rodent species recorded from the British Pleistocene are 
then discussed in systematic order and, finally, an attempt is made to relate a generalized sequence 
of rodent faunas to the climatic sequence usually employed today as the basis for British 
Pleistocene chronology. 



Nearly half a century has elapsed since the publication by the British Museum 
(Natural History) of M. A. C. Hinton's Monograph of the Voles and Lemmings in 
1926. Today it is still the only comprehensive work on British Pleistocene rodents 
and remains the standard reference work on this subject. In the meantime there 
have been great advances in many related fields of study. Hinton did not attempt 
to relate his sequence of rodent faunas to the glacial-interglacial sequence of climatic 
events, which was already gaining favour at the time of his studies, and which is 
known today to be even more complicated than the four glacial and three interglacial 
stages for so long employed as a basis of Pleistocene chronology. 

Excavations in the British Isles since 1926 have produced a wealth of new fossil 
rodent material which was not available to Hinton. Findings from the Westbury- 
sub-Mendip Fissure (Somerset), Swanscombe (Kent), Tornewton Cave (Devon) and 
a series of Late Glacial cave sites in the Peak District (Staffordshire and Derbyshire), 
excavated and published by various workers, are of special importance. The advent 
of carbon-14 (^^C) dating has permitted the accurate dating of some Upper Pleisto- 
cene rodent remains. Studies of pollen and insects are further, relatively new, 
stratigraphic aids. 

On the continent of Europe there have been great advances in rodent studies 
which provide a basis for the interpretation of rodent migrations into the British 
Isles and assist in identifying stages of geographical isolation, W. von Koenigswald's 
recent work (1972, 1973) on the Mimomys-Arvicola lineage, based initially on 
continental European remains and subsequently extended to the British Isles, 
throws new light on the British Pleistocene rodent sequence. 

In the present work the authors have cooperated on a reappraisal of the Pleistocene 
rodent faunas of the British Isles. One of us (K. Kowalski of Krakow, Poland) is 
mainly responsible for the systematic part and for comparison with continental 
evidence ; the other (A. J. Sutcliffe of London) for the stratigraphic sequence of 
British rodent faunas. Discussions with W. von Koenigswald (Tubingen) about the 
Mimomys-Arvicola lineage have contributed fundamentally to the last-mentioned 
part of this work. 

Although we cover ^y species of rodents from over 100 sites, many problems re- 
main which can only be resolved by further careful collecting. Additional material 
is needed from critical deposits such as the early and late parts of the Forest Bed 
Series of Norfolk and from Hinton's Middle Terraces of the Thames, or from deposits 
of equivalent age. Some of the faunal assemblages which we have listed, such as 
that from the Otter Stratum of Tornewton Cave, are unique and cannot be compared 
with deposits of the same age in other parts of the British Isles. We have shown 
scarcely any evidence of differences between contemporary faunas in different areas, 
nor of temporal faunal changes within the broad climatic phases used for reference. 
The systematic relationship between some of the rodent species is still far from clear. 
All these aspects of rodent studies and many others offer great scope for future 
research. If this Bulletin provides a stepping stone for such work it will have been 
worth while. 


A. History of Studies 

Although publications of the eighteenth century and, in greater number, of the 
first decades of the nineteenth contain references to the occurrence of remains of 
beaver in British Quaternary deposits, the first scientific description of fossil rodents 
from this country did not appear until the middle of the nineteenth century. In 
1846 R. Owen published a detailed description, accompanied by excellent drawings, 
of the remains of Trogontherium cuvieri and Castor fiber from the Forest Bed deposits 
of East Angha. He also mentioned the presence of the remains of Arvicola 'which I 
have been unable satisfactorily to distinguish from Arvicola amphibia, or common 
Water-rat' (1846 : 202) in the sediments of many caves. The bones of the same 
species were present, according to Owen 'in newer pliocene deposits'. He considered 
(1846 : 205) the specimens from the 'older pliocene crag near Norwich . . . indicated 
a species of Arvicola intermediate in size between the Water-vole {Arvicola amphibia) 
and the Field-vole [Arvicola arvalis)' . 

The next important contribution to the study of British Pleistocene rodents was 
that of W. A. Sanford. His papers on rodents from caves of Somerset (1870a, b) 
contain not only descriptions of forms still living in Britain {Arvicola terrestris, 
Clethrionomys glareolus and Microtus agrestis) but also of species now extinct in this 
country {Microtus oeconomus, Lemmus 'of the type of L. norvegicus' and Dicrostonyx 
torguatus). It is interesting to note that Sanford correctly determined the skulls of 
D. torquatus but ascribed jaws of the same species to Arvicola gulielmi. He men- 
tioned also the occurrence of a small hamster, 'Cricetus songarus (Pallas)', and 
Spermophilus erythrogenoides , a species previously discovered and described by H. 
Falconer (in Murchison 1868). 

The first attempt to make a systematic classification of fossil voles, one of the 
most difficult groups of rodents, was made by H. P. Blackmore & E. R. Alston 
(1874). They failed to recognize the difference between Recent and fossil species, 
but were the first to record Microtus nivalis in Britain. 

A series of works on the 'preglacial' deposits of East Anglia by E. T. Newton 
(1881, 1882a, b, 1890b, 1891 and 1909) provides an important contribution to the 
knowledge of the rodents of this area. Newton was the first to state that the vole 
remains were specifically different from Recent forms and he named a new species, 
Arvicola intermedius Newton 1881. He also published diagnoses of two other new 
species of rodents, a hamster, 'Cricetus vulgaris Runtonensis n. subsp.' and a beaver 
'Trogontherium minus n. sp.' His work is of importance in regarding the vertebrate 
remains, including many rodents, from the 'pre-glacial' deposits of East Anglia as 
specifically distinct from living forms. He also published the first scientific descrip- 
tion (1894, 1899a, b) of the vertebrate remains, including rodents, from the rich 
late Pleistocene fauna of Ightham Fissures, Kent. 

The greatest contribution to the study of British Pleistocene rodents was that of 
M. A. C. Hinton (1883-1961). His first note on this subject appeared in 1900 and 
later this group of mammals became the main object of his interest. His work 
culminated in 1926 with the publication of volume i of his Monograph of the Voles 
and Lemmings. Hinton soon became the world authority on Microtinae and his 
monograph (unfortunately unfinished) is still the standard work in this branch of 


mammalogy. He was the first to recognize that, since rodent species have short 
ranges in time, their remains are of special stratigraphic value in the study of the 
Pleistocene. They thus help the geologist attempting to correlate scattered or 
isolated cavern deposits with others to which ordinary stratigraphical methods can be 
applied. The importance of this concept was further expanded by Kowalski (1966). 
Hinton (1926a ; 1926b : 126-136) recognized a series of rodent faunas of different 
ages at British localities. In order of increasing antiquity these were as follows : 

Third Terrace of the Thames, in the valley of the River Lea. 

Ightham Fissure (Kent) stage. 

Late Middle Terrace of the Thames (typical locality Crayf ord and Erith, Kent) . 

Early Middle Terrace of the Thames (typical locality Grays Thurrock, Essex) . 

High Terrace of the Thames (Ingress Vale, Swanscombe, Kent). 

Upper Freshwater Bed at West Runton, Norfolk. 

Shelly Crag at East Runton, Norfolk. 

Norwich Crag and Weybourne Crag of the Norfolk Coast. 

Although Hinton did not attempt to relate tl is division to any sequence of climatic 
fluctuations, his faunal stages were far ahead of any other palaeontological division 
of the British Pleistocene at that time. 

Hinton also laid the basis for systematic study of the Microtinae (Savage 1963). 
After 1926 he abandoned nearly completely his study of the British fossil rodents, 
being involved in problems of pest control and the study of other mammals. He 
nevertheless published in 1952 a thorough description of rodent remains from the 
Late Glacial deposits of the Lea Valley. 

Hinton's work is not only an important chapter in the history of zoology, but it 
still forms the standard reference on British Pleistocene rodents. For this reason it 
seems necessary to mention some of its limitations. He was a monoglacialist, or 
strictly an antiglacialist, as he postulated only one very late and only moderately 
cold period in the Pleistocene history of England. This opinion, which he regarded 
as the established truth, probably also influenced his zoological views. When he 
came across the fossil remains of species associated today with arctic climate he 
was inclined to determine them as forms different from Recent ones, since they 
could not then be used as evidence of particular climatic conditions. 

Hinton was a prominent typologist. Although he was not more of a 'splitter' 
than most of his contemporary zoologists, he nevertheless used subspecific names 
extensively and was inclined to determine each new variant as a new species. For 
this reason, even in Hinton's own collection, many intermediate specimens were left 

Finally, Hinton accepted the multituberculate origin of rodents, a theory now 
generally rejected. He accepted that microtine evolution has always led from 
forms with complicated to forms with simple teeth. Strangely enough this did not 
hinder him in the correct reconstruction of some of the lines of evolution in voles, 
since among the different tendencies in the late evolution of the teeth of this par- 
ticular group some lines did indeed progress from more complicated to simpler 


Relatively few contributions to the study of British Pleistocene rodents have 
been published since the appearance of Hinton's monograph. These include the 
work of J. W. Jackson (1932, 1934) ; L. S. Palmer (1934) ; A. Schreuder (1929, 
1931, 1950, 1951) ; J. N. Carreck (1957, 1966) ; D. Bramwell (i960, 1964, 1970) ; 
A. J. Sutcliffe & F. E. Zeuner (1962) ; J. C. Pernetta (1966) ; R. J. G. Savage 
(1966) ; K. Kowalski (1967) ; W. von Koenigswald (1973) and M. Bishop (1974, 
1975). Most of these papers contain site lists of rodents with relatively little general 
discussion of their stratigraphical or systematic position. The papers by Schreuder 
(1929, 1931, 1951) are worthy of special mention for containing a thorough re- 
description and systematic discussion of the beaver remains from Britain. The 
paper of 1950 contains a list of the voles from the stratigraphically important 
Middle Pleistocene deposits of Swanscombe, Kent. More recently A. J. Stuart, in a 
general review of the British Pleistocene fauna (1974), listed the rodent faunas from 
key sites and discussed their stratigraphical occurrence. 

Since the last synthetic work of Hinton (1926b) there has been great progress in 
other fields of study related to the problem of the history of British rodents. Investi- 
gations on the continent of Europe (not without influence from Hinton's work) have 
brought to light many rich local rodent faunas of Pliocene and Pleistocene age and 
have made it possible to reconstruct in greater detail the stratigraphical position 
and evolutionary history of many of these mammals. Recent contributions in the 
field include the work of J. Chaline (1972), who gives a detailed description of the 
rodents of the Middle and Upper Pleistocene of France, and of W. von Koenigswald, 
who studied the phylogenetic lineage of the genus Arvicola. In addition, knowledge 
of the systematic position, ecology and geographical variability of Recent rodents 
has greatly improved throughout the whole of the Palaearctic region. 

In the British Isles there have been extensive geological and palaeobotanical 
studies (e.g. West 1968), which have provided a more detailed stratigraphic back- 
ground for rodent studies. In addition the stratigraphic position of many old 
localities has been clarified and new rodent localities have been discovered. A 
new aspect of investigation has been provided by zoological studies of insular races 
of small mammals (Corbet 1961). 

In view of the increasing importance of rodents in stratigraphic studies and of all 
the additional information which has become available since the publication of 
Hinton's monograph, it is timely to review the existing data on British Pleistocene 
rodents. In the description which follows the stratigraphic age of crucial rodent 
localities of the British Isles will be further examined, the systematic position and 
synonymy of the rodent species, currently overburdened with too many names 
connected with insufficiently characterized forms, will be discussed and the history 
of the rodent population of Britain during the Pleistocene reviewed. Many gaps 
nevertheless remain in the known sequence of events. This paper does not contain 
morphological descriptions of the various species of British Pleistocene rodents, a 
task, important for further stratigraphic and systematic studies, which needs to be 
carried out in the future. In the meanwhile subspecific designations are not used 
in the present paper. The nomenclature of living species of rodents here used is 
based mainly on the work of Ellerman & Morrison-Scott (1966). Information about 


morphology, biology and distribution of Recent rodents in Britain is available in 
Miller (1912), Matthews (1952), Southern (1964) and Corbet (1966). Beirne (1947) 
has discussed the possible arrival dates of some species of rodents into the British 

B. The Geological Background 

As previously mentioned, great progress has been made towards a more detailed 
understanding of the British Pleistocene sequence since Hinton published his mono- 
graph in 1926. Some deposits considered to be Pliocene at that time are now 
regarded as Lower Pleistocene, and there now exists a detailed picture, still being 
further elaborated from current studies, of alternating glacial and interglacial 
stages in the British Isles. A recent contribution of special importance is that of 
West & Wilson (1966), who demonstrated from palaeobotanical studies that the 
Cromer Forest Bed Series of Norfolk in fact represents two interglacial stages with 
an intervening cold stage. 

At the present time the most generally accepted correlation for the British 
Pleistocene is that recommended by the Geological Society of London (Mitchell et al. 
1973), which is shown, with some additional information from other sources indicated, 
in Table i. 

While in general it seems possible to relate most British rodent faunas to this 
sequence, there are some parts (notably the Cromerian and Wolstonian-Ipswichian 
stages) which seem inadequate to account for all the rodent stages currently 
attributed to them. There can be little doubt that they are more complex than is 
indicated in the table opposite. 

Although, in the 1973 correlation of the Geological Society, the Wolstonian and 
Ipswichian are recognized as two stages only, an earlier correlation published by the 
same Society (Evans 1971) proposed a more detailed chronology for this part of 
Pleistocene time which must be mentioned here, since it has important application 
to our rodent studies. 

From a consideration of information derived from deep-sea cores and other lines 
of evidence Evans argued that there had been more than one warm phase since the 
Hoxnian (Holsteinian) Interglacial. Whilst accepting the period from about 100 
to 70 thousand years ago (which he regarded as the true Last Interglacial, the 
Ipswichian or Eemian, equivalent to Zeuner's Late Monastirian shoreline) as the 
only fully warm period of considerable length since the Holsteinian, he drew atten- 
tion to other lesser mild phases of post-Holsteinian age about 170 000 and 130 000 
years ago, possibly equivalent to the Danish Vejlby I and Vejlby II mild stages, the 
possible relationship of which is shown in Table 2. 

Evans also drew attention to some problems related to the interpretation of this 
part of the Pleistocene sequence. He pointed out that Zeuner, who regarded the 
Last Interglacial as double, used the term 'Last Interglacial' for two different ranges 
of time, that lasting from 180 to 120 thousand years ago (cycles 5w and 4) and also 
for that lasting from 130 to 70 thousand years ago (cycles 4W and 3). He considered 
that some confusion had arisen in archaeological circles where 'Last Interglacial' 
had been used as a reference datum and preferred to restrict this term only to zone 3W. 



Generalized British Pleistocene sequence 





















"S c 



^ 2 















u « u 

u H °a < Q 


&:; H 

W en « 

S W „ S 

O £ R « 

« O W w 

O tin W C/} 



z o 

u 1 




^^5 ; 

J. 1 




2 + 


2 + 




5 ^ 

^ < 

S o 


Z H 

g I 


n ^ 








c .2 



ri tn 


C/) W 











O &- 






























1 2 
1 ^ 


O J 

^. ^> 



Table 2 

Possible relationship of continental climatic divisions to British mild phases (after Evans 1971) 

Evans' half-cycles Years ^ Continental Mediterranean 

(c = cold, w = warm) ago ° climatic divisions sea levels 

^ / o names 

2C 70 000 Early Weichsel, Early Wiirm 

3w 100 000 Ipswichian Eemian Late Monastirian 

3c 120 000 Saale 2, Riss 2, Warthe 

4w 130 000 Vejlby 2 Main Monastirian 

4c Minor cold phase 

5w 180 000 Vejlby i, ? Domnitz 

5c 200 000 Saale i, Riss i 

6w HoxNiAN Holsteinian 

It follows from Evans' chronology that the terms Saale 2, Riss 2 and Penultimate 
Glaciation of some authors have been used for periods of time later than Last Inter- 
glacial of some other authors, and that great confusion can arise here if stage names 
such as these are applied incautiously to our studies of British Pleistocene rodents. 
Likewise the terms Penultimate Glaciation or Wolstonian (when applied away from 
the type locality) could confusingly be used to mean any period of time from that 
considered by some writers to be the middle of the Hoxnian/Holsteinian (for ex- 
ample, Mullender's suggestion in Wymer (1974) that the upper part of Lower Loam 
of Swanscombe is Wolstonian) to that immediately preceding Evans' Interglacial 
half-cycle 3W. Wymer (1974) has suggested a warm stage during the Wolstonian. 
Bristow & Cox (in Mitchell et al. 1973) have argued, from their study of glacial 
deposits in East Anglia, that the interglacial deposits at the Hoxnian type locality 
of Hoxne belong to the last and not penultimate interglacial and they referred the 
Ipswichian and Hoxnian deposits, which they accepted may have been separated 
by a cold oscillation, to a single interglacial between the Devensian and Anglian. 

In the present paper we will try to overcome such problems, as far as possible, 
by concentrating on establishing a relative chronology for the rodent faunas of the 
British Isles which will not be affected by future refinements in the naming of the 
British Pleistocene sequence, rather than to attempt to refer these faunas too rigidly 
to a chronology which is clearly incomplete. Let us now make a detailed examina- 
tion of the rodent faunas from the various British Pleistocene localities. 


Remains of Pleistocene rodents have been found at many localities in the British 
Isles, notably in the early Pleistocene marine Crag and Forest Bed deposits of East 
Anglia and in later river terrace and cave deposits. In Ireland they are known only 
from cave deposits of late Pleistocene age. The location of the principal localities 
is shown in Fig. i. 

A. Deposits of East Anglia 

Extensive areas of Suffolk and Norfolk are covered by Crag and Forest Bed 
deposits from which rodent remains have sometimes been recovered. 


Although recent research (West & Wilson 1966, Norton 1967, West 1968) has 
greatly increased understanding of the early Pleistocene sequence of East Anglia, 
all the rodent remains from this region available for the present study are unfor- 
tunately from old collections, many of them with imprecise stratigraphic information. 
A series of rodent faunas can nevertheless be distinguished. 

(i) Red Crag. This is a marine shallow- water shore deposit typically laid down 
in land-locked bays. It is best developed in Suffolk where it consists mainly of 
shelly sands. There is also a basal nodule bed vs^ith rolled and polished fossils, 
including mastodon teeth, apparently derived by the Crag sea from earlier deposits. 
The occurrence of these derived fossils, which include mammalian remains of Eocene, 
Miocene and Pliocene age, makes the study of the Red Crag mammalian fauna 
extremely difficult. For a long time the deposit was considered to be of Pliocene 
age and it was not until 1948 that its Lower Pleistocene date was accepted and the 
occurrence of relatively unmineralized contemporary mammalian remains, associated 
with the derived fossils, was recognized. 

Only a few rodent remains have been found in the Red Crag. Trogontherium 
minus (known from Astian localities on the European continent) and Hystrix, 
represented by teeth (Spencer 1966) and by gnawing on part of a deer antler (Sutcliffe 
& CoUings 1972), are probably derived from pre-Pleistocene deposits and cannot be 
included in the contemporary Red Crag faunal list. 

Castor fiber, recorded from a number of Red Crag localities, including Sutton and 
Woodbridge (47, Fig. i) is probably a contemporary species. A rolled microtine 
tooth from the Red Crag has been provisionally referred by Spencer (1964) to 
Mimomys sp. 

(ii) IcENiAN Crag. A further series of marine deposits, the Icenian Crag (in- 
cluding the Norwich and Weybourne Crags), occupies a basin on the north of the 
Red Crag outcrop. It was apparently laid down in a more open sea. Its exact 
relationship to the Red Crag is not fully understood, but its age appears to range 
from Thurnian, through Antian and Baventian, to Pastonian. According to West 
(1968), these stages may be equivalent to the Tiglian, Eburonian and Waalian in 
the stratigraphy of the Netherlands. The relationship between the Norwich and 
Weybourne Crags is also not clear. The Weybourne Crag, typified by the mollusc 
Macoma halthica, is found at more than one stage. The youngest part of the 
sequence, including part of the Weybourne Crag, is of Pastonian age and is con- 
temporaneous with the lowest part of the Cromer Forest Bed series. Part of the 
Norwich Crag of Suffolk may also be Pastonian. 

Rodent remains, which have been found in the Norwich and Weybourne Crags of 
Bramerton (55, Fig. i), Covehithe (51), Easton Bavents (50), Sizewell (49), Thorpe 
(48), Trimingham (61) and other locaHties represent Mimomys pliocaenicus , M. 
reidi, M. newtoni, Castor fiber and Trogontherium boisvilletti. Since conditions were 
unsuitable for the preservation of very small remains, it is not surprising that no 
teeth of gHrids and murids have been found in the deposits. M. pliocaenicus and 
M. reidi are typical Tiglian (Upper Villafranchian) elements and confirm a Lower 
Pleistocene age for at least part of this series. 



Fig. I. Location map of the principal Quaternary rodent localities in the British Isles. 
Key to numbers opposite. 





Joint Mitnor Cave, 
Buckfastleigh, Devon 


Marlow, Buckingham- 



Levaton Cave, Torbryan, 


Isleworth, Middlesex 




Cray ford and Frith, Kent 


Tornewton Cave, 
Torbryan, Devon 


Ightham Fissures, 
Ightham, Kent 



Brixham Cave, Brixham, 


Northfieet, Kent 




Swanscombe, Kent 


Happaway Cave, 


Upnor, Kent 

Torquay, Devon 


Murston, Kent 


Kent's Cavern, Torquay, 


Grays Thurrock, Essex 

71 (ii) 



Aveley, Essex 

7 (i) 

Cow Cave, Chudleigh, 


Ilford, Essex 


36 (i) 

Hackney, London 

71 (iii) 


Chudleigh Fissure, Devon 

36 (ii) 

Angel Road, Middlesex 


Huntspill Cut, Huntspill, 

(north London) 


36 (iii 

) Ponders End, Middlesex 



Brean Down, nr Brean, 


Nazeing, Lea Valley, 


Uphill Cave, Uphill, 


Water Hall Farm, 



Hay Wood Rockshelter, 


Hitchin, Hertfordshire 


Hutton, Somerset 



lO (ii) 

Bleadon Cave, Bleadon, 





Cambridge Fens 

lo (in 

) Picken's Hole, 

(including Burwell and 

nr Bleadon, Somerset 

Swaffley), Cambridge- 


lo (iv 

) Hutton Cave, Hutton, 




Copford, nr Colchester, 


Banwell Cave, Somerset 



II (i) 

Rowberrow Cavern, 


Clacton, Essex 

Burrington, Somerset 

44 (i) 

Harkstead, Suffolk 


II (ii) 

Aveline's Hole, 

44 (ii) 

Stutton, Suffolk 

Burrington, Somerset 


Felixstowe, Suffolk 



Gough's Cave, Cheddar, 

46 (i) 

Bobbitshole, nr Ipswich, 


Fissure, Somerset 

46 (ii) 

Stoke Tunnel Beds, 
Ipswich, Suffolk 



Clevedon Cave, Clevedon, 

47 (i) 

Sutton, Suffolk 


47 (ii! 

) Woodbridge, Suffolk 



Alveston Fissure, Alveston 

47 (iii 

i) Kyson, Suffolk 



Thorpe, Norfolk 


16 (i) 

Minchin Hole, nr Penard, 

(see p. 96) 



Sizewell, Suffolk 


16 (ii) 

Bacon Hole, nr Penard, 


Easton Bavents, Suffolk 



Covehithe, Suffolk 

17 (i) 

King Arthur's Cave, 


Kessin gland, Suffolk 


nr Whitchurch, 


Geldeston, nr Beccles, 



17 (ii) 

Great Doward Cave, 


Hoxne, nr Diss, Suffolk 




Bramerton, Norfolk 



Happisburgh, Norfolk 



Merlin's Cave, Symond's 


Ostend, Norfolk 

Yat, Herefordshire 


Bacton, Norfolk 




Paston, Norfolk 




Mundesley, Norfolk 


Upton Warren, 


Trimingham, Norfolk 


nr Droitwich, 


Overstrand, Norfolk 



Cromer, Norfolk 



Sugworth, Oxfordshire 


East Runton, Norfolk 


Fisherton, Salisbury, 


West Runton, Norfolk 



Swanton Morley, Norfolk 



Thatcham, Berkshire 


Lynx Cave, Denbighshire 

24 (i) 

West Wittering, Sussex 

(exact location not 

24 (11) Selsey, Sussex 



Gwaenysgor Cave, 

Prestatyn, Flintshire 
Elder Bush Cave, 

Wetton, Staffordshire 
Harborough Cave, 

Brassington, Derbyshire 
Fox Hole Cave, High 

Wheeldon Hill, Earl 

Sterndale, nr Buxton, 

Etches' Cave, Dowel Dale, 

Earl Sterndale, nr 

Buxton, Derbyshire 
Dowel Cave, Dowel Dale, 

Earl Sterndale, nr 

Buxton, Derbj^shire 
Langwith Cave, Upper 

Langwith, nr Scarcliffe, 

Pin Hole Cave, Creswell, 

Hessle, nr Kingston- 

upon-HuU, Yorkshire 
Staple Howe, nr 


Star Carr, nr 


Kirkdale Cave, Helmsley, 

Cowside Cave No. 3, 

Settle, Yorkshire 
Dog Holes Cave, 

Warton Crag, nr 

Carnforth, Lancashire 
Middlestots Bog, Edrom 

Parish, Berwickshire, 

Corstorphine, nr 

Edinburgh, Midlothian 
Loch of Marlee, Kinlock, 

Creag nan Uamh Cave, 


Keshcorran Caves, 

Ballymote, Co. Sligo, 

Republic of Ireland 
Edenvale Caves, nr Ennis, 

Co. Clare 
Red Cellar Cave, nr 

Lough Gur, 

Co. Limerick 
Castlepook Cave, 

Doneraile, Co. Cork 
Castletownroche Cave, 

Connaberry, Co. Cork 
Kilgreany Cave, 


Co. Waterford 
Ballynamintra Cave, 

Whitechurch, Co. 

Nornour, Isles of Scilly 





♦ Overstrand 
H/p, ^^Mundesley 




jThorpe <P 


Easton Bavents 

Weybourne Crag & Forest Bed • 
Norwich Crag ▲ 
Red Crag ♦ 
Fig. 2. Location map of Crag and Forest Bed rodent localities in East Anglia. 

(iii) Cromer Forest Bed Series. This differs from the earlier Crag deposits in 
being predominantly estuarine and freshwater, with some beach deposits. Coastal 
exposures occur at many localities along the Norfolk coast, though these are un- 
fortunately becoming increasingly obscured by sea defence work. Abundant 
remains of mammals (including rodents) and of plants occur in the Forest Bed. The 
most important rodent localities are West and East Runton (65, 64), with fewer 
remains from Ostend (57), Cromer (63), Overstrand (62), Mundesley (60), Paston 
(59), Bacton (58) and Kessingland (52). 

It has long been recognized that all the Forest Bed deposits are not contem- 
poraneous but that they accumulated over a considerable period of time, during 
which there occurred changes of climate and fauna. West & Wilson (1966), basing 


their conclusions on a study of plant remains, identified four climatic stages within 
the Forest Bed Series of deposits. In descending order these are : 

Early Anglian (cold, followed by glacial conditions 

during the later Anglian) 
Cromerian sensti stricto (warm) 
Beestonian (cold) 

Pastonian (warm) 

Rodent remains are known from all horizons except the Beestonian. The Ostend 
Forest Bed presents a problem which is further discussed on pp. 48, 122. 

(a) Pastonian of East Runton and Happishurgh. Only the earliest part of the 
Forest Bed sequence is represented at East Runton (64) ; deposits of Cromerian age, 
sensii stricto, apparently being absent. Azzaroli (1953) observed that the large 
mammals from this locality are Villafranchian species and that mammals present 
in the later part of the Forest Bed are absent. West (personal communication, 
1972) considers, from palaeobotanical studies, that no Cromerian deposits, sensu 
stricto, have yet been proved on the foreshore at East Runton. He found, in 
descending order, the following sequence : 

c. Clay Conglomerate, being reworked estuarine sediment, probably of Pastonian 
age, redeposited during late Pastonian or Beestonian times. 

b. Shelly Crag, of Pastonian age, regarded by Hinton (1926b : 365) as Weybourne 

a. Flint Bed, relating to a pre-Pastonian land surface. 

Unfortunately the East Runton section has not been seen really well for many 
years and all the rodent remains available for the present study are from old collec- 
tions. Three species of Mimomys are recorded ; these are M. newtoni, M. plio- 
caenicus and M. savini. The occurrence, together in the Shelly Crag, of the last 
two species, which represent successive stages in the development of the same 
phyletic line of voles, suggests the natural or accidental mixing of elements from 
different layers. M. pliocaenicus and M. newtoni are represented in both the Shelly 
Crag and among the 'clay pebbles' of East Runton, the holotype of M. newtoni being 
from the former horizon. 

The precise stratigraphic range of these three species is far from clear. M. savini 
(represented by a few specimens from the Shelly Crag) is very abundant in the later, 
Cromerian sensu stricto, deposits of West Runton. The occurrence of remains of 
M. pliocaenicus and M. newtoni (known also from the earlier Norwich Crag) in the 
'clay pebbles' could mean that these rodents persisted into Pastonian times. Such 
an interpretation would not be out of harmony with the 'Villafranchian' megafauna 
of this locality. The rolled condition of some of the remains could alternatively 
indicate their derivation from an earlier deposit. 

Trogontherium boisvilletti and Castor fiber have also been found at East Runton. 

A second Forest Bed locality, where the deposits are apparently of Pastonian 
age only, is Happisburgh (56). Remains of Castor fiber and also fir cones bearing 


marks which appear to indicate gnawing by squirrels (Newton 1882a) have been 
found there. 

(b) Beestonian. No rodent remains are known from Beestonian deposits. 

(c) Cromerian sensu stricto. The Upper Freshwater Bed of West Runton (65), 
defined by West (1961) as the type deposit of the Cromerian interglacial, contains 
abundant rodent remains. The following species are represented : Apodemus 
sylvaticus, Trogontherium boisvilletti, Castor fiber, Cricetus cricetus, Mimomys savini, 
Clethrionomys glareolus, Pitymys arvaloides, P. gregaloides, Microtus arvalinus, M. 
nivaloides and M. raUicepoides. Muscardinus recorded by Hinton (in Barrett- 
Hamilton & Hinton 1910-21, 2 : 351) from the 'Forest Bed' can possibly be added 
to the above list. 

The rodent fauna with at least 11 species is unusually rich. It is predominantly 
of forest and meadow type, testifying to a mild climate ; no arctic elements are 
present. The presence of Cricetus in this fauna is difficult to explain. 

Sciurus whitei is represented from the marine 'Monkey Gravel' at West Runton, 
which overlies the Upper Freshwater Bed. 

(d) Anglian. Newton (1882b) described some isolated worn teeth of Spermophilus, 
found in association with remains of arctic plants, in the Arctic Freshwater Bed 
near Mundesley (60). This deposit, which is immediately overlain by glacial till, 
has been interpreted as an indication of oncoming cold conditions in early Anglian 

The rodent fauna of the Forest Bed deposits at Ostend (57) near Bacton presents 
a stratigraphic problem, since it cannot be accurately related to the above sequence. 
The remains include Arvicola hactonensis and A. greeni (Hinton 1926b : 386, 389), 
now both regarded as synonyms of ^. cantiana, which led Hinton to conclude (p. 391) 
that the Ostend deposit is later than the Upper Freshwater Bed of West Runton. 
The specimens were collected over a century ago by the Rev. C. Green and un- 
fortunately lack reliable stratigraphic information. 

B. Terrace and Solifiuxion deposits of the Rivers Thames and Lea 

The richest series of rodent-bearing deposits related to a British river system is 
that of the River Thames and River Lea. The deposits of these rivers have been 
extensively worked commercially for gravel and brickearth and there have been 
many deep excavations for building foundations in the London area, leading to 
frequent discoveries of rodent remains (Fig. 3). 

Three main series of deposits occur in association with these rivers, the exact 
relationship between which is often difficult to determine. Firstly, there are the 
terrace deposits of their upper courses, including Marlow and Isleworth in the Upper 
Thames, and Water Hall Farm Pit, Nazeing, Ponders End, Edmonton and Hackney 
on the River Lea. Deposits of both interglacial and glacial age occur here, those 
representing glacial stages merging with buried channels in the Lower Thames 
Valley province, where they are inaccessible for study in consequence of later rise 
of sea level. 


Fig. 3. Location map of rodent localities in the Thames estuary and the Lea valley. 

Secondly, there are terraces of the Lower Thames, including Swanscombe, 
Clacton, Ilford, Aveley and Grays, Crayford and Erith. Hinton (1926b : 126- 131) 
classified these as the High Terrace of the Thames (Swanscombe), the Early Middle 
Terrace (Grays Thurrock), and the Late Middle Terrace (Crayford and Erith). 
Whereas each terrace of the upper part of the river is of approximately constant 
height above the present-day bed of the river and thus becomes progressively lower 
as it is followed downstream, these terraces of the Lower Thames (which are related 
to former high sea levels) have approximately horizontal aggradation surfaces 
and they do not extend further upstream than the head of the contemporary tidal 
limit of the river. All are of interglacial or interstadial age and they can be cor- 
related with raised beaches of similar heights along the open coastline. The cold 
stages which occurred between the accumulation of these terraces of the Lower 
Thames are represented by deposits submerged in the buried channels of the present- 
day river, previously mentioned. 

Thirdly there are solifluxion and hillwash deposits, the heights of which are un- 
related to the terrace system, with the deposits of which they are often interbedded. 
Northfleet is the only rodent locality, probably of this category, known to the 

It follows that, in an area as complicated as the valleys of the Rivers Thames and 
Lea, the exact relationship between the various rodent-bearing deposits is difficult 
to interpret. The literature on this subject, which goes back for nearly a century 
and a half, is voluminous. The general sequence was discussed by King & Oakley 
(1936) and in a number of important papers by Zeuner (e.g. 1954), who produced a 
schematic section from the Ebbsfleet Valley to Swanscombe, Kent, showing his 
interpretation of the relationship between the various deposits there. He inter- 
preted the solifluxion and loess deposits with Levallois industries of Baker's Hole 


and Ebbsfleet as earlier than deposits of both Main Monastirian and Late Monastirian 
age, which he regarded as Last Interglacial, although only the Late Monastirian is 
accepted as Last Interglacial by Evans (p. 40). 

During recent years J. N. Carreck has made a detailed further study of many of 
the mammalian localities in the Thames estuary, especially those in Kent, together 
with related museum collections and previous literature. On a basis of all lines of 
evidence he relates {in litt.) the deposits of some of the more important localities, 
here given in ascending stratigraphic order, as follows : 

1. Interglacial deposits of Swanscombe, with Clactonian and Acheulean industries. 

2. Interglacial deposits of Grays Thurrock and Little Thurrock (including the 
Orsett Road brickearths, but not those of West Thurrock, which are slightly 
later), Ilford and Aveley. The Ilford mammoth is a transitional form between 
Mammuthus irogontherii and M. primigenius. The Ilford fauna does not 
include any cold elements. 

3. Deposits of the Baker's Hole cold stage at Ebbsfleet/Northfleet. The first 
true mammoth, M. primigenius, intermediate in form between the mammoths 
of Ilford and Crayford, appears at this locality in non-estuarine deposits over- 
lying coombe rock (solifiuxion) deposits. There are several associated Leval- 
lois industries. 

4. Interglacial deposits of Crayford and Erith. Carreck found that the mammoth 
from the Lower Crayford Brickearth is more advanced than that from stage 3, 
though not so advanced as Last Glaciation mammoths from such localities as 
Ponders End, in the Lea valley, north London. He suggested that a layer of 
shattered chalk at the base of the Crayford sequence may be equivalent to the 
Baker's Hole cold stage and that the Crayford Gravel and Lower Brickearth, 
which are interglacial, follow immediately in time. The Crayford fauna 
suggests more open grassland conditions than indicated at Ilford or Grays, and 
there is a Levallois industry more advanced than that at Baker's Hole. 

5. Other interglacial deposits, with hippopotamus, suggest that there may have 
been a further amelioration of climate. Carreck considers that the hippopo- 
tamus may have survived to a very late stage during the Last Interglacial. 
The Last Glaciation was characterized by a more advanced mammoth than 
that from Crayford. 

Carreck pointed out that most of the Grays mammals were collected over a century 
ago, possibly from more than one terrace, which makes the study of this locality 
difficult ; he drew attention to the hazards of confusing the Baker's Hole cold stage, 
which he regarded as occurring within the Ipswichian Interglacial, with the Wol- 
stonian/Saale Glaciation. 

Carreck's chronological conclusions are of very far-reaching importance, with 
two points of special interest. Hinton's supposition that the Crayford deposits are 
later than those at Grays receives further support, but unlike the Geological Society 
(Mitchell et al. 1973), which regards the Northfieet coombe rock as Wolstonian and 
earlier than Ilford (which is regarded as Ipswichian), Carreck regards the Ilford 
deposits as being earlier than those of Northfieet. 


Let US now consider in more detail the various deposits in the valley of the Thames 
and its tributaries where remains of fossil rodents have been found. 

(i) Lower Thames. 

(a) High Terrace of Swanscomhe. One of the most important British rodent 
localities is the world-famous Swanscombe skull site (see Ovey (1964) for a fuller 
account). A series of estuarine deposits is aggraded to a height of approximately 
31-5 m (103 ft) O.D. and is believed to have accumulated when the sea level rose to 
this height. Solifiuxion deposits bring this figure to a total of 35*5 m (113 ft). 
Rodent remains have been found in two gravel pits, separated by a distance of about 
a third of a kilometre and now both disused. These are Barnfield Pit, where skull 
fragments of Acheulean Man were found, and Dierden's Pit, Ingress Vale. The 
following sequence of deposits has been described from Barnfield Pit : 

6. The Upper Gravel. This is regarded as a solifiuxion deposit, later than the 
terrace proper. No mammalian remains have been found in it. 

5. The Upper Loam, with an Acheulean industry but no mammalian remains. 

4. The Upper Middle Gravel, occupying a channel in the underlying deposits and 
containing an Acheulean industry. The top of this stratum, which is at about 
31-5 m (103 ft) O.D., is regarded as the true surface of the terrace deposits 
proper. This deposit contains the 'Homo layer' in which the human remains 
were found. 

3. The Lower Middle Gravel, with an Acheulean industry. 

2. The Lower Loam, with an in situ Clactonian activity horizon at its base and a 
knapping floor higher up. A zone of subaerial weathering, with many well- 
preserved animal footprints, is present on the surface of this deposit. 

I. The Lower Gravel, with an early Clactonian industry, resting on a bench of 
underlying rock at 21-27 m (70-90 ft). 

The deposits contain a rich moUuscan fauna which has recently been re-examined 
by Kerney (1971). He concluded that the Lower Gravel, Lower Loam and the very 
base of the Lower Middle Gravel accumulated under temperate conditions, after 
which the climate became cooler. 

Rodent remains have been found in the Lower Gravel (Carreck 1959), in the Lower 
Loam (remains recently excavated by Waechter), and in a silt bed in the Upper 
Middle Gravel at a level slightly higher than the 'Homo layer' (Schreuder 1950). A 
few rodent remains have also been found at Ingress Vale. The distribution of rodent 
species at these two localities is shown in Table 3. 

The exact relationship between the deposits of Barnfield Pit and Ingress Vale is 
uncertain. Arvicola cantiana occurs at both the latter site and in the Lower Loam 
of Barnfield Pit, suggesting a possible correlation of the two. Kerney (1971 and 
personal communication) found at Ingress Vale a temperate molluscan fauna associ- 
ated with an industry which is culturally more advanced than that in the Lower 
Gravel and Lower Loam, suggesting to him a possible equivalence to the base of the 
Lower Middle Gravel. 

The Swanscombe rodent remains are insufficiently abundant to permit reconstruc- 
tion of any environmental and climatic changes, but the species of voles found in the 



Table 3 

Distribution of rodent species at Barnfield Pit and Ingress Vale, Swanscombe, Kent 

Barnfield Pit 





Trogontherium boisvilletti 

Castor fiber 


Apodemus sylvaticus* 

Clethrionomys glareolus 

Arvicola cantiana 


Lemmus sp. 

Microtus arvalinus 


M. ratticepoides 


Microtus sp. {arvalis-agrestis group) 

Pitymys arvaloides 



* Cited in earlier papers as A . whitei. 









Lower Loam suggest that during the deposition of this layer a meadow environment 
may have prevailed. There are no arctic and no forest species, but such negative 
evidence is not decisive. 

Although the Swanscombe deposits are generally regarded as being of Hoxnian 
Interglacial age, there is increasing evidence that a substantial interval may have 
elapsed between the deposition of the Lower Gravels/Lower Loam complex and of 
the overlying Middle Gravels, In general the rodent fauna of the lower unit quite 
closely resembles that of the Upper Freshwater Bed of West Runton. Pitymys 
arvaloides, Microtus arvalinus and M. ratticepoides occur in both the Upper Fresh- 
water Bed at West Runton and in the Lower Loam at Barnfield Pit. The two above- 
mentioned species of Microtus persist into the Upper Middle Gravel. Mimomys 
savini, characteristic of Cromerian assemblages sensu stricto, is nevertheless absent 
from Swanscombe, where it is replaced by Arvicola, absent from the Upper Fresh- 
water Bed. It is of interest that by Upper Middle Gravel times Lemmus had 
appeared and A. cantiana may have disappeared. At the present day Lemmus is a 
rodent of northern latitudes. Whereas we have no means of demonstrating that its 
habit is not a recent adaptation, the occurrence of this genus could be interpreted 
as further evidence of the cool conditions indicated by the mollusca. 

Recently Mullender (in Wymer 1974) has examined pollen from the Lower Loam 
of Barnfield Pit and found a marked break in the profile with a great increase in pine 
and near disappearance of alder. He equated the lower part of the Lower Loam 
with the Hoxnian Late-temperate Zone II and the upper part with the Wolstonian. 
If this interpretation were to be accepted then the later deposits of Barnfield Pit 
could no longer be referred to the Hoxnian, which would lead us to fresh problems 
of nomenclature for this part of the sequence. 

(b) Clacton. The exact age relationship between the deposits of Clacton and 
Swanscombe is not clear, though Clacton is generally regarded as a further Hoxnian 


locality intermediate in age between the Lower Loam and Middle Gravel of Barnfield 
Pit. The Clacton deposits, which lie at about sea level, are at a lower altitude than 
those at Swanscombe. West (1972) suggests that slight downwarping has occurred 
in the Clacton area, which may explain this difference. There is a Clactonian in- 
dustry. Plant remains from Clacton, studied by Pike & Godwin (1952), suggest 
about one-third of an interglacial sequence, during which mixed deciduous forest of 
the warmth maximum was replaced by coniferous forest. 

Only a few rodent remains have been found in the Clacton deposits. These are 
Trogontherium cf. cuvieri (= T. hoisvilletti) and Clethrionomys sp. (Singer et al. 1973), 
Castor sp. and Microtis of the agrestis group (Hinton 1923b), and Arvicola cf. 
praeceptor {= A. cantiana) (Warren 1955, quoting Hinton). 

(c) Early Middle Terrace at Aveley, Grays Thurrock and Ilford. Hinton (1926b : 
129) defined Grays Thurrock (33) as the type locality of this terrace of the Thames, 
with which the present writers also include Ilford (35) and Aveley (34). Table 4 
lists those rodent species whose remains have been found at these localities. 

The age of the deposits at these localities has been the subject of much discussion 
which is yet to be concluded. Hinton (1926b : 129- 131) considered that the 
species of Arvicola from Grays {A. praeceptor, = A. cantiana) was not closely related 
to modern species of that genus. He considered that the latest Forest Bed deposits 
(presumably Ostend), the High Terrace of the Thames and the Early Middle Terrace 
were close in time. Zeuner (1945) considered Grays to be Hoxnian. 

Hoxnian terrace deposits do indeed occur in the Grays area, though remains of 
fossil mammals do not seem to have been found in them. Wymer (1957) records 
terrace deposits of Hoxnian age with Clactonian implements resting on a bench at 
15 m (49 ft) O.D., but points out that the Grays brickearth is later than this deposit. 
Most of the Grays rodent remains were found in the brickearth in a small pit near 
Orsett Road (Hinton 1901), about 650 m (700 yds) west of Wymer's site. 

West (1969) studied plant remains from the three sites mentioned above and 
concluded that Aveley and Ilford are of Ipswichian age, and that Grays is inter- 
glacial, probably also Ipswichian. He pointed out that there was at this time 
apparently an important aggradation phase which resulted in the spreading of 
alluvium up to levels of between 12 and 15 m (40-50 ft) O.D. 

The age of the Grays-Ilford-Aveley deposits will be further discussed below. 
From the evidence of the Arvicola remains, Grays Thurrock would appear to be 
relatively early. 

Table 4 

Distribution of rodent species at Grays Thurrock, Aveley and Ilford, Essex 

Grays Thurrock Aveley Ilford 

Castor fiber x x 

Apodemus sylvaticus x 

Clethrionomys glareolus X X 

Arvicola cantiana XXX 

Microtus agrestis X Microtus sp. X 


(d) Late Middle Terrace at Crayford and Erith. A rich rodent fauna has been 
collected from the Thames terrace deposits of Crayford and nearby Erith ; it has 
been described by Kennard (1944). Owing to the lenticular character of the deposits 
at these localities there is little constancy in the details of the succession. Three 
broad divisions can, however, be identified. In descending order there are : 

3. The Upper Brickearth, apparently not fluviatile but the result of sludging. 
2. The Lower Brickearth and Corhicula Bed, up to 9 m (30 ft) O.D., laid down 

in a sluggish stream and in more strongly running water respectively. Leval- 

lois artefacts have been found in this deposit. Most of the rodent remains 

are from the Corhicula Bed. 
I. The Lower Gravel, deposited by a fast-flowing river. The top was a land 

surface occupied by Levallois man, who left many artefacts. 

Most of the rodent remains have been found in the Corhicula Bed, which is a 
sandy development of the Lower Brickearth. The following species are represented : 
Spermophilus primigenius, Microtus oeconomus, M. nivalis, M. agrestis, Arvicola sp., 
Lemmus lemrmts and Dicrostonyx torquatus. 

The age of the Crayford deposits is a topic requiring extensive further study. 
Hinton (1926b : 131) considered Crayford to be later than Grays, with a major 
intervening change of fauna. This view is also supported by Carreck {in litt.). At 
the present time Crayford is widely considered to be of Ipswichian age and both the 
freshwater and land moUusca found in the Corhicula Bed suggest conditions warmer 
than at the present time. The associated rodent fauna, however, is a typical 
assemblage of the 'penultimate' Glaciation. Spermophilus primigenius is unknown 
from the sediments of the Last Glaciation in Europe and, although Microtus nivalis 
was present during the Last Glaciation in many parts of Europe, and still survives in 
the mountains of central and south Europe, there is no evidence that it survived 
into the Last Glaciation in Britain. 

It has been suggested that the lemmings burrowed into the Corhicula Bed from a 
later land surface, or alternatively that the remains were derived from an earlier 
deposit. Field evidence, however, suggests that the rodent remains are contem- 
porary with the Corhicula Bed. 

A total absence of forest species of land mollusca indicates that the country was 
open grassland. Living species of Spermophilus are also predominantly grassland 
animals. This could be interpreted as the beginning or end of an interglacial stage, 
though it is pointed out by Turner (in prep.) that riverside trampling by large 
herbivores can also give rise to clearances in otherwise wooded areas. 

The Crayford deposits apparently date from some time during Wolstonian- 
Ipswichian times. Their precise stratigraphic position will be further discussed 
below and in Section IV (pp. 125-126). 

(e) Floodplain Terrace complex. A series of lower terraces of Last Interglacial 
and Last Glaciation age in the Thames estuary have produced an abundance of 
remains of large mammals but, with one exception [Castor from the Upper Flood- 
plain Terrace of the River Medway, a river flowing into the Thames estuary, at 
Upnor, Kent), no rodent remains are known from them. 


(f ) The Middle Terrace jFloodplain Terrace problem. Although no stratigraphically 
significant rodent remains have been found in the Floodplain Terrace complex 
of the Thames estuary, and although Hinton did not mention this terrace in his 
monograph (his 'Third Terrace of the Thames' is actually a terrace of the upper part 
of the River Lea) , abundant rodent remains which are probably of Upper Floodplain 
Terrace age have nevertheless been found in many British cave deposits. The 
present section would be incomplete without some discussion of the status of this 

It has already been pointed out that various workers have presented evidence 
that the period of time regarded by the Geological Society (Mitchell et al. 1973) as 
Hoxnian-Wolstonian-Ipswichian was, in fact, more complicated than this sequence 
suggests. Zeuner (1945) described evidence for a minor cool phase dividing what 
he regarded as the Last (Ipswichian) Interglacial into an earlier and a later part. 
He considered that there were two stages of high sea level during the Last Inter- 
glacial, the Main Monastirian or 18 m and subsequent Late Monastirian or 8 m 
shorelines, interrupted by an intra-Monastirian fall of sea level. These stages were 
accepted by Evans (1971 ; see pp. 40-42), though he argued that the earlier of these 
two stages should not be called Last Interglacial and observed that some confusion 
seemed to have arisen in archaeological circles where 'Last Interglacial' had been 
used for both. 

Zeuner correlated the so-called Taplow Terrace of the Thames estuary (our 
Middle Terrace complex belongs here ; the term 'Taplow' is best abandoned in the 
estuary since Taplow is in the upper part of the Thames and the terrace there is not 
estuarine) with the Main Monastirian sea level and the Upper Floodplain Terrace 
with that of the Late Monastirian. Both these supposed terraces are highly fossili- 
ferous, with apparently different mammalian megafaunas. 

When Zeuner described his interpretation of the Last Interglacial terrace sequence 
of the Thames estuary very little palaeobotanical evidence was available to him. 
During recent years, however, fossil plant remains have been found at a number of 
Last Interglacial locahties there, the most important being Trafalgar Square 
(Franks i960). Seven Kings Station, Ilford (West et al. 1964) and Aveley (West 
1969), and here problems have arisen. The plant remains from these localities 
were assigned to the pollen zones shown in Table 5. 

To those who had been studying the terraces and mammahan faunas of the 
Thames estuary, these results were surprising. Whereas Ilford and Aveley had 
appeared to be part of the Middle Terrace of the Thames and Trafalgar Square part 
of a lower terrace, the Upper Floodplain Terrace (i.e. Zeuner's two terraces of the 
Last Interglacial, representing two distinct periods of time), only one climatic 
fluctuation, with the climatic optimum of Zone lib represented at all three sites, 
could be recognized from the pollen evidence. Was Zeuner wrong in separating the 
two stages of his supposed Last Interglacial, the Middle and 'Upper Floodplain' 
Terraces of the Thames estuary really being only one terrace, or are the floral 
remains from two separate climatic events so similar that these terraces cannot be 
distinguished on palaeobotanical evidence? Mammahan and morphological evi- 
dence suggests that, in agreement with the views of Zeuner and Evans, the latter 

56 pleistocene rodents 

Table 5 

Pollen zonation of some 'Ipswichian' interglacial sites in the Thames estuary 

Pollen Zones 


Ilford Aveley 

Trafalgar Square 

Last Glaciation 



Pine, birch 




Oak, hornbeam, 
silver fir 

^^ , Mammoth 





Mixed oak forest 

^^^ ^^^ ' elephant 






Birch, pine 



N.B. The Trafalgar Square deposits are of Zone lib age, and should be shown somewhat higher than 

alternative is the more likely even though palaeobotanical evidence supports only a 
single terrace (Fig. 4). 

Even the chronological interpretation of the Ilford sequence is not without 
problems. West et al. (1964) referred the gravel under the Ipswichian brickearth 
and plant deposit at Seven Kings Station to the Gipping cold stage, but Carreck 
{in litt.) has pointed out that there is considerable lateral variation of the Ilford 
deposits with interglacial species of moUusca and mammals frequently present in the 
gravels and sands. 

Let us now consider the large mammals of these Thames sites in some detail. At 
Trafalgar Square (supposedly Upper Floodplain Terrace) Zone lib contains remains 
of hippopotamus, straight-tusked elephant, a rhinoceros which is probably Dicero- 
rhinus hemitoechus, fallow deer, red deer, giant ox, bison, lion and other animals. 
No mammoth or horse remains were found. This is a faunal assemblage which 
occurs commonly in British cave deposits with localities as widely spread as Joint 
Mitnor Cave, Devon, and Kirkdale Cave, Yorkshire (Sutcliffe i960). 

The Ilford-Aveley fauna appears to be entirely different. At Ilford most of the 
elephant remains are of an early form of mammoth with affinities to the Middle 
Pleistocene Mammuthus trogontherii, though straight-tusked elephant is also 
represented. Two species of rhinoceros are present, D. hemitoechus and D. kirch- 
bergensis. This last species is also common in the preceding Hoxnian Interglacial. 
There is an abundance of horse ; both hippopotamus and fallow deer are absent. 



At Aveley straight-tusked elephant in Zone lib is replaced by mammoth in Zone III. 
Comparison of rodent species unfortunately cannot be made, since no diagnostic 
remains have been found in the Upper Floodplain Terrace. Arvicola cantiana is 
present in the Middle Terrace at Aveley, Ilford and Grays Thurrock. 

The megafaunas of Aveley and Ilford on the one hand, and of Trafalgar Square 
on the other, are so different that it is difficult to believe that they are contemporary, 
though consideration must be given to the possibility of their representing different 
stages within a single climatic fluctuation. 

The evidence does not seem to support the latter alternative. The hippopotamus 
level at Trafalgar Square is Zone lib. A vast amount of Zone lib clay was exca- 
vated for cement-making at Aveley but no hippopotamus remains were found. The 
Ilford plant remains cover Zones I A- lib but there were no hippopotamus remains 
there ; two specimens formerly attributed to this animal have since been shown to 
have been incorrectly identified. 

It has been pointed out, in support of there having been only one temperate stage, 
that most of the Ilford mammal remains were recovered over a century ago, during 
excavations for brickearth which were relatively shallow and did not extend into 


SCHEME A (Morphological & Mammal Evidence) 

SCHEME B ( F^laeobotanical Evidence) 

Ilford Mammals 

Trafalgar Square Mammals 

Fig. 4. Schematic section of Thames estuary Ipswichian terraces, showing alternative 
interpretations of the stratigraphic succession. Reproduced from Sutchffe & Bowen 
(1973 : 18). 


underlying gravel deposits, which were of no commercial value. The organic 
deposit described by West et al. (1964) at Seven Kings Station underlay the brick- 
earth, so it is possible that the Ilford mammalian fauna is later than Zone lib 
(perhaps Zone III), that is the Trafalgar Square fauna might have been found in the 
basal part of the Ilford terrace had excavations been carried deeper. According to 
this theory Trafalgar Square represents an earlier stage than Ilford ; according to 
the twin terrace theory it is later. The former interpretation presents several 
difficulties. In spite of extensive commercial excavation of sand and clay in the 
London area, no such relationship has ever been observed ; the Upper Floodplain 
Terrace does appear to be a good morphological feature ; the Ilford mammoth is 
relatively primitive ; the Ilford fauna has never been recognized in any British cave, 
with the possible exception of Hutton Cave, whereas the Trafalgar Square fauna 
with hippopotamus occurs frequently, suggesting that such deposits have been 
subject to less denudation and are more recent. 

In the absence of diagnostic rodent species in the Upper Floodplain Terrace of the 
Thames the above problem cannot at present be resolved from studies in the Thames 
area. It will be further critically examined below, in the section on hippopotamus 
faunas in caves and in Section IV. 

(ii) Upper Thames. 

(a) Isleworth. Remains of Microtus oeconomus and M. gregalis were found at 
Willment's Gravel Pit, Isleworth, in a deposit where they were associated with 
remains of temperate mollusca and insects (Coope, 1975) and with plant remains 
dated by ^*C to 43 140 + 1520 or — 1280 years b.p. (Birmingham 319). This deposit 
was overlain by sands and gravels containing remains of reindeer and cut by many 
fossil ice wedges. A description of the site, by J. Simons, is in preparation. The 
sequence is interpreted as indicating deteriorating climatic conditions after an inter- 
stadial during the Last Glaciation. It is probably the earliest occurrence of M. 
gregalis in Britain ; this species appears to be restricted there to the Last Glaciation 
and to have disappeared before the end of that stage. 

(b) Mar low. A small sample of brickearth, found in the Treacher collection at 
the British Museum (Natural History) and labelled as coming from a brickearth pit 
I mile north-east of Marlow, was found to be rich in rodent remains, predominantly 
dissociated teeth. They were examined by Dr G. B. Corbet who found that most of 
them were apparently of Microtus arvalis. There was also one first lower molar of 
M. oeconomus. The lack of any second upper molars of M. agrestis and the occur- 
rence of only one first lower molar of M. oeconomus among the many first lower 
molars (which teeth distinguish these last two closely related species from M. 
arvalis) suggest M. arvalis was the principal species present. 

The age of the Marlow rodents is uncertain, but it is likely to be Last Glaciation. 
Neither M. arvalis nor M. oeconomus survives on the mainland of Britain at the 
present day. 

(iii) River Lea. 

(a) Water Hall Farm Gravel Pit. An important sequence of Pleistocene deposits 
occurs at Water Hall Farm, Hertfordshire. Here the valley of the River Lea has 


cut through earlier glacial deposits which still cap the hills on either side. In the 
valley bottom the River Lea is flanked on one bank by its present-day floodplain, 
on the other by a slightly higher terrace in which have been found abundant remains 
of interglacial mammals, including Hippopotamus and Palaeoloxodon antiquus. 
Remains of Mammuthus have also been found. At the base of the sequence of 
deposits in this terrace there was formerly exposed a white marl with remains of 
amphibia and of the rodents Microtus agrestis, M. oeconomus and M. nivalis. Since 
most of the commercial excavation of the interglacial terrace, since concluded, was 
carried out by mechanical excavators, the exact stratigraphic relationship between the 
hippopotamus layer and the rodent layer remains unproved, but nearly the whole of the 
ossiferous gravel deposit for which the pit was worked overlay the rodent-bearing 
marl and there seems no doubt that the rodent remains antedate those of the hippo- 
potamus. Unfortunately no rodent remains have been found in the upper deposits. 
The sequence of events at this locality is provisionally determined as follows : 

1. Boulder clay and glacial deposits. Not later than 'penultimate' glaciation. 

2. Retreat of ice and valley deepening by River Lea. 

3. Deposition of marl with remains of amphibia and rodents. 

4. Deposition of interglacial deposits with Hippopotamus and P. antiquus over- 
lying the rodent marl. Last Interglacial. 

5. Further deepening of Lea Valley and formation of present floodplain. Last 
Glaciation to present day. 

(b) Nazeing, Ponders End, Edmonton and Hackney. A complex series of deposits, 
mainly of late Last Glaciation and Holocene age, occupies the valley of the lower 
part of the River Lea. Rodent remains have been found at a number of localities 
there. The most important of these is at Nazeing, where a series of channel, lake 
and marsh deposits have been palaeobotanically dated as ranging from Late Glacial 
to post-Glacial vegetation Zone VII, providing the latest known survival dates in 
Britain for a number of rodent species (Hinton 1952). The late Pleistocene part 
of the sequence produced remains of Microtus oeconomus, M. gregalis, Arvicola 
terrestris, Lemmus lemmus and Dicrostonyx torquatus. Of these the last-mentioned 
species and M. gregalis were still present in Late Glacial pollen Zone III and M. 
oeconomus appears to have survived possibly until post-Glacial Zone V. Arvicola 
terrestris also persisted into the Flandrian and Apodemus sylvaticus, Clethrionomys 
glareolus and M. agrestis appear in Zones V-VI, by which time post-Glacial re- 
afforestation had reached an advanced stage. 

Remains of Dicrostonyx torquatus have been found at three other late Pleistocene 
localities in the Lea Valley : Ponders End (Warren 1916), Angel Road, Edmonton 
(Hinton 1912) and at Hackney. 


(a) Northfleet, Ehbsfleet and Baker's Hole. Burchell (1935) recorded finding rodent 
remains associated with those of mammoth and rhinoceros in non-estuarine deposits 
fining a channel cut through a coombe rock (solifluxion) deposit at Baker's Hole, 
between Northfleet and Swanscombe, Kent. As previously mentioned (p. 50) 
Carreck {in litt.), from the study of the mammoth remains from this site, considered 


that the deposits, which indicate an amehorating climate following the deposition 
of the coombe rock, are later than those of Ilford but earlier than Crayford. Stuart 
{in litt.), who has recently re-examined the Burchell specimens in the British Museum 
(Natural History) together with a few specimens collected more recently by Kerney 
and Sieveking, gives the following list of species : Clethrionomys glareolus, Microtus 
arvalisjagrestis and Arvicola cantiana. He found that in teeth of the last-mentioned 
species the enamel is clearly differentiated, as in Mimomys, with no A. cantiana- 
terrestris intermediate forms (see notes on Arvicola, pp. 99-102). This suggests a 
relatively early date for the Northfleet rodent remains, a conclusion in agreement 
with Carreck's observations on the mammoth remains. Burchell (1935) listed 
Microtus arvalis (here interpreted as M. arvalisjagrestis group), M. nivalis and M. 
anglicus (= M. grcgalis) from his excavations, but the last two have not been 
confirmed by Stuart's re-examination of the available specimens. 

C. Cave Deposits 

Remains of Pleistocene rodents have been found in many caves in the British 
Isles. Most such deposits accumulated in consequence of animals accidentally 
falling down shafts or of their remains being carried into caves by birds of prey. 
Since, however, the entrance parts of caves are vulnerable to destruction by processes 
of denudation, many formerly-existing rodent deposits have since disappeared ; it is 
unusual to find any rodent remains in caves of earlier than Upper Pleistocene age. 
Kent's Cavern and a recently discovered fissure at Westbury-sub-Mendip, Somerset, 
are probably the only exceptions. A Lower Pleistocene cave deposit with megafauna 
has also been found at Dove Holes, Derbyshire (Crag age), but unfortunately no 
contemporary rodent remains have been recorded. In contrast, cave deposits with 
Last Glaciation and Holocene rodent remains are very numerous. 

Few of these sites have produced stratified sequences of rodent faunas, either 
because the deposits accumulated during only a short period of time, because they 
had been disturbed before excavation, or because they were excavated before the 
importance of stratigraphy had become adequately appreciated. 

The following are the most important rodent caves in the British Isles. 

(i) Westbury Fissure (13). The recently discovered fissure infilling at Westbury- 
sub-Mendip, described by Bishop (1974, 1975), is of special importance, since its 
rich mammalian fauna indicates a stage slightly later than the type Cromerian of 
West Runton, not otherwise well represented in the British Pleistocene. Bishop 
described, in descending order, the following sequence of deposits : 

III. 'Rodent Earth' (layer 10, the upper part of the 'Calcareous Group', layers 
2-9 below). A deposit with an abundance of remains of rodents and other 
small mammals, possibly an accumulation of pellets of birds of prey. 
II. 'Calcareous Group' (layers 2-9, excluding the 'Rodent Earth'). Pre- 
dominantly limestone breccias with abundant remains of bears. The cave 
was probably a bear den at this stage. 
I. 'Siliceous Group' (layer i). Water-laid deposits with some rolled bones and 



Table 6 

Stratigraphic distribution of rodent species in the Westbury Fissure 
(after Bishop 1974, wi1h personally communicated additions) 


IIL 'Rodent Earth' 

II. 'Calcareous Group' 
'Rodent Earth') 

I. 'Siliceous Group' 



The distribution of rodent species in the Westbury fissure is shown in Table 6. 

The age of the deposit has been discussed in detail by Bishop (1974). From the 
'Calcareous Group', excluding the 'Rodent Earth', he recorded a fauna including 
Homotherium latidens, Felis gombaszoegensis, Ursus deningeri, Xenocyon lycaonoides, 
Canis lupus mosbachensis , Dicerorhinus etruscus and Equus mosbachensis. Rodents 
from this level are Apodemus sylvaticus, Lemfmis sp. (the earliest record of this genus 
in the British Pleistocene), Arvicola cantiana, Pitymys gregaloides and Microtus 
arvalinus. The same species are represented in greater abundance in the 'Rodent 
Earth', which Bishop regarded as the last stage of the 'Calcareous Group'. Remains 
of Pliomys episcopalis (the first record in the British Pleistocene), Dicrostonyx sp. 
(the earliest record in the British Pleistocene) and Clethrionomys were also found in 
the 'Rodent Earth'. 

Bishop regarded the Westbury fauna as later than the type Cromerian of West 
Runton but not later than Elsterian. He drew attention to its similarity to that of 
the classic 'late Cromerian' sites of Europe (in particular Mauer, Hundsheim, Tarko 
and Mosbach) and equated the Westbury 'Rodent Earth' with the Arvicola fauna 
group I of Koenigswald (1973) (see pp. loo-ioi). This stage was previously 
unrecognized in the British Pleistocene sequence. 

(ii) Kent's Cavern. Unfortunately most of the rodent remains found in Kent's 
Cavern were excavated during the nineteenth century and only sparse stratigraphic 
information is associated with them. The following species have been found there 
(Hinton 1915, Kennard 1945-6, British Museum (Natural History) collection) : 


Castor fiber, Apodemus fiavicollis, Clethrionomys glareolus, Arvicola terrestris, Dicro- 
stonyx torquatus, Lemmus lemmus, Pitymys gregaloides, Microtus agrestis, M. 
oeconomus and M. gregalis. In addition, Campbell & Sampson (1971) recorded a 
specimen of Arvicola greeni (regarded here as a synonym of A. cantiana) among 
specimens collected from the cave by J. MacEnery between 1825 and 1829. This is 
a typically Last Glaciation-Holocene fauna, with the exception of P. gregaloides, 
not recorded elsewhere in Britain in deposits later than those of the Westbury 
Fissure, and A. cantiana. The occurrence of P. gregaloides and also sabre-toothed 
cat, Homotherium, in Kent's Cavern has been interpreted by both Hinton (1926b) 
and Campbell & Sampson (1971) as evidence of a Cromerian stratum somewhere in 
Kent's Cavern. 

The sequence of deposits in the cave varies from one chamber to another (for 
details see Campbell & Sampson). The principal deposits, in descending order, are 
as follows : 

5. Black Mould, with Mesolithic and later artefacts and fauna. 

4. Granular Stalagmite, with Mesolithic and Neolithic artefacts and fauna. 

3. Cave Earth, including a local area of hearths known as the Black Band. Middle 
to Upper Palaeolithic industries. Mammalian fauna including woolly mam- 
moth, woolly rhinoceros and hyaena. 

2. Crystalline Stalagmite. 

I. Breccia, with many bear remains and with a Lower Palaeolithic industry. 

The upper part of the above sequence is typically Last Glaciation (the Cave 
Earth) to Holocene. Most of the rodent remains probably come from the upper 
levels. Pitymys gregaloides, on the other hand, is a Middle Pleistocene species 
probably derived from the Breccia, the age of which is at present uncertain. Homo- 
therium, P. gregaloides and A. cantiana are all recorded from the Westbury Fissure. 
At the present time it seems most likely that the earliest remains from Kent's 
Cavern are of Westbury Fissure age. There is no conclusive evidence of fauna as 
early as Cromerian sensu stricto. It may be inferred that the Crystalline Stalagmite 
represents a major break in sedimentation, possibly because the cave entrance had 
become sealed. 

(iii) ToRNEWTON Cave (3). The most important sequence of stratified cave 
deposits with rodent remains known from the British Isles is that of Tornewton Cave, 
south Devon (Sutcliffe & Zeuner 1962, Kowalski 1967). Deposits in the shaft-like 
Main Chamber of this cave and of the talus deposits outside span a cold-interglacial- 
cold sequence which is unique in showing differences between the rodent faunas of 
the two cold stages concerned. The relationship between this sequence and that of 
the terraces of the River Thames will be considered in Section IV, pp. 124-127. 

The principal deposits in the Main Chamber of Tornewton Cave (excluding some 
superficial deposits excavated during the nineteenth century and a series of stalag- 
mite floors), in descending order, were found to be as follows : 

5. 'Diluvium'. Most of this deposit was excavated during the nineteenth century. 
The remaining part contained a few remains of wolf, hyaena, bear, bovid and 


reindeer. A Holocene moUuscan fauna has been identified by M. P. Kerney 
from this horizon and it is probable that the mammalian remains are a mixture 
of derived Pleistocene and contemporary Holocene species. 
4. The Reindeer Stratum. Most of this deposit was excavated from the Main 
Chamber of the cave during the nineteenth century. Abundant rodent 
remains, associated with remains of wolf, hyaena, horse, rhinoceros, reindeer, 
a bovid and a sparse Upper Palaeolithic industry were nevertheless recovered 
more recently from an extension of this stratum in the talus outside the cave. 
The stratum is of Last Glaciation age. 
3. The Hyaena Stratum. A deposit with abundant remains of spotted hyaenas, 
which animals apparently occupied the cave as a lair, associated with occasional 
remains of hippopotamus, narrow-nosed rhinoceros, red and fallow deer, lion, 
wolf and bear. This is an interglacial assemblage, tentatively referred to the 
2. The Bear Stratum. A deposit with abundant remains of brown bears, which 

animals used the cave as a lair at this stage. 

I. The Glutton Stratum. The earliest fossiliferous deposit in the cave, with 

numerous remains of brown bears associated with occasional remains of glutton 

(wolverine) and reindeer. This deposit represents an earlier cold phase. 

Table 7 shows the stratigraphic distribution of rodent remains in the Main 

Chamber and (Reindeer Stratum only) in the talus of Tornewton Cave. It will be 

seen from the table that, in the Glutton Stratum, Microtus oeconomus and M. 

nivalis make up the bulk of the rodent fauna. Lagurus lagurus (from the only 

British locality with this genus), Allocricetus bursae and Cricetus cricetus are typical 

species of the 'penultimate' glaciation. Dicrostonyx torquatus and Lemmus lemmus 

Table 7 
Number of rodent specimens from the main deposits of Tornewton Cave 

Glutton Bear Hyaena Reindeer ,^., . , r^ , ^ 

o^ ^ o^ ^ o^ X c^ ^ Diluvium Total 

Stratum Stratum Stratum Stratum 

Apodemus sylvaticus 






Cricetus cricetus 






cf. Allocricetus bursae 






Dicrostonyx torquatus 






Lemmus lemmus 





Clethrionomys glareolus 






Arvicola sp. 






Microtus agrestis 






Microtus nivalis 






Microtus oeconomus 






1 109 

Microtus gregalis 







Lagurus lagurus 







Total 1660 31 9 218 113 2031 

The count is based on the number of determinable specimens, not individuals. The apparent greater 
abundance of remains in the Glutton Stratum is the result of more extensive sampling of this deposit. 
Remains from minor deposits not included above bring the total number of specimens determined to 


are present but not abundant. Microtus gregalis is absent. Sylvan species are very 
rare, though a few remains of Clethrionomys glareolus occur with this predominantly 
meadow and steppe assemblage. A single tooth of Apodemus sylvaticus probably 
represents contamination of the deposit. The rodent fauna of the Glutton Stratum 
has affinities to that of Crayford, Kent, from which it was probably not greatly 
separated in time. 

Only a few rodent remains were collected from the Bear Stratum. The fauna 
does not differ in composition from that of the underlying Glutton Stratum though, 
with the exception of one tooth of Lagunis lagurus and four of Microtus nivalis, 
the typical 'penultimate glaciation' elements are lacking. Their absence could be 
accidental, however, since they are also rare in the Glutton Stratum. The absence 
of lemmings could indicate a slight amelioration of climate. 

Rodents are poorly represented in the Hyaena Stratum. Apodemus sylvaticus, 
represented by only one specimen, is a forest species. Microtus agrestis and Arvicola 
sp. are ecologically neutral. 

The Reindeer Stratum has a predominantly cold fauna with the typically Last 
Glaciation Microtus gregalis and with Dicrostonyx torquatus. 

The apparent mixture of arctic and sylvan species in the 'Diluvium' is in agree- 
ment with this deposit being of Holocene age, with derived Pleistocene remains. 
Apodemus sylvaticus reappears here. The climate was milder than before. 

The sequence of rodent faunas described above is of great importance as it makes 
possible, for the first time, distinction between mammalian assemblages, indis- 
tinguishable on the basis of the larger mammals, of the two cold stages concerned. 
Arctic elements (for example, lemmings) appear twice in the sequence, but are lack- 
ing from the interglacial Hyaena Stratum. The lower cold stage is characterized 
by, among other rodents, Cricetus cricetus, cf. Allocricetus hursae, Lagurvts lagurus 
and Microtus nivalis, all absent from the Reindeer Stratum, where they are replaced 
by large quantities of M. gregalis which, on the other hand, is absent from the 
Glutton Stratum. 

Distinction is also possible between the teeth of Arvicola from the Glutton Stratum 
and those from the Reindeer Stratum. W. von Koenigswald (1973 and personal 
communication) has found that although the form from the latter deposit is A. 
terrestris, that from the Glutton Stratum is an intermediate form tending towards 
Arvicola cantiana, known from the Middle Pleistocene deposits of Swanscombe (see 
pp. 100-102). 

In addition to the deposits of the Main Chamber of Tornewton Cave two important 
rodent deposits were found in positions where they could not be directly related to 
the main sequence. These are the Otter Stratum, in a small chamber adjoining the 
Main Chamber, and the Upper Rodent Stratum, on the rock platform outside the 
cave mouth. 

The Otter Stratum was found to contain an abundance of remains of brown bears 
associated with some mammalian species unrecorded elsewhere in the Pleistocene of 
the British Isles, notably Cyrnaonyx, the clawless otter, and Crocidura, the white- 
toothed shrew (Rzebik 1968). It was composed of a mixture of broken stalagmite 


blocks, some containing faunal remains, in an earthy matrix, suggesting some dis- 
turbance of the deposit. The following rodent specimens were collected from it, 

Cricetus cricetus 2 specimens 

Lagurus lagurus 1 specimen 

Dicrostonyx torquatus i specimen 

Lemmus lemmus i specimen 

Microtus nivalis 4 specimens 

Microtus oeconomus 82 specimens 

Apodemus sylvaticus 2 specimens 

Clethrionomys glareolus 19 specimens 

Microtus agrestis 5 specimens 

Arvicola sp, 8 specimens 

This fauna appears to be a mixture from two originally separate layers. Remains 
from a warm period, including Microtus oeconomus, predominate. They point to a 
last interglacial age. No Hoxnian elements are present. Lagitrus lagurus, Cricetus 
cricetus and Microtus nivalis, on the other hand, are species of the 'penultimate' 
glaciation. In an attempt to separate the two faunal assemblages some of the 
stalagmite blocks were dissolved in acid. Only temperate species {Microtus 
agrestis, i determinable specimen ; M. oeconomus, 6 ; Arvicola sp,, i ; Clethrionomys 
glareolus, 4 ; Apodemus sylvaticus, i) were found in the stalagmite. The exact 
stratigraphic relationship between the Otter Stratum and the deposits of the Main 
Chamber of Tornewton Cave is unfortunately uncertain, since the overlying deposits 
were excavated during the nineteenth century and are no longer available for 
examination. The cave was apparently still inhabited as a lair by bears, making it 
probable that the deposit antedates the Hyaena Stratum. On the other hand, it 
seems to be later than the Glutton Stratum since Koenigswald has found that, 
whilst the Arvicola from both the Glutton Stratum and the Otter Stratum is an 
intermediate form between A . cantiana and A . terrestris, that from the Otter Stratum 
is slightly more advanced, though not so advanced as the form from the Reindeer 
Stratum, which is A. terrestris. The Otter Stratum fauna appears to be a unique 
assemblage, unknown from any other locality in the British Isles, 

The deposit known as the Upper Rodent Stratum occupied a rift on the rock 
platform outside the main entrance to Tornewton Cave. It was only a short distance 
below the surface and is probably of modern date. The following species are 
represented : Apodemus sylvaticus, 58 specimens ; Clethrionomys glareolus, 14 ; 
Microtus agrestis, 86 ; Arvicola terrestris, 2 ; Micromys minutus, i. All these 
species live near the cave at the present day, no extinct species are represented ; and 
M. minutus is a late arrival probably associated with neighbouring plough-land. 

In conclusion the relative proportions of the various ecologic groups of rodents 
represented in Tornewton Cave are shown in Table 8. 

(iv) Other caves with pre-Tpswichian' deposits. In light of the faunal 
sequence demonstrated at Tornewton Cave, it would be surprising if other rodent 
faunas dating from the time interval between the Hoxnian and the warm stage 
represented by the Hyaena Stratum of Tornewton Cave were not to be found in 
caves from time to time. At present this period is one of some obscurity. 


Table 8 

The relative proportions 

of the various ecologic 


of rodents found 

in Tornewton C 


and steppe 




Upper Rodent Stratum 










Reindeer Stratum 





Hyaena Stratum 





Bear Stratum 





Glutton Stratum 





Tundra and steppe elements include Dicrostonyx torquatus, Lemmus lemmus, Microtus gregalis, M. 
nivalis, Lagurus lagurus, Cricetus cricetus and Allocricetus. Boreal elements include M. oeconomus. At 
the present day this species lives mainly in the taiga belt but it also occurs, under milder climatic con- 
ditions, in marshland. The forest element is represented by Apodemns sylvaticus and Clethrionomys 
glareolus. Arvicola sp. and M. agrestis are 'neutral' in their ecologic requirements. Micromys has been 
omitted from the table since it was probably introduced by man. 

Some isolated pre-Tpswichian' rodent-bearing cave deposits have nevertheless 
apparently been found. Hinton (1926b) regarded the rodents of Clevedon Cave (14) 
and Banwell Cave (10), Somerset, as probably contemporary with those of Crayford 
which, we have observed, are similar to those of the Glutton Stratum of Tornewton 
Cave. The rich material from Clevedon Cave is of special interest, as it contains 
only two species of rodents, Microtus nivalis and M. oeconomus. Hinton (1907a) 
wrote : 'Dr H. C. Male very generously presented me with a small series of the 
numerous jaws of Microtus which he found in the Clevedon deposit.' In the material 
described by Hinton, which is still preserved in the British Museum (Natural 
History), there are exclusively remains of the species mentioned above. It is pos- 
sible that Dr Male did not present Hinton with the material of the other rodents, 
but it does not seem possible that he retained jaws of Microtus agrestis, as they are 
difhcult to distinguish from those of M. nivalis except by a specialist. It seems 
probable that the Clevedon Cave remains accumulated at a time when the rodent 
fauna was composed exclusively of M. oeconomus and M. nivalis. 

Hinton's specimens from Banwell Cave have not been seen by the present writers. 
The only available specimens, representing Arvicola sp. and Microtus sp., are 
stratigraphically inconclusive. 

The Gough's Cave (12) rodent fauna is composed of Arvicola sp., Microtus agrestis, 
M. oeconomus and M. nivalis. The last-mentioned species suggests an age equivalent 
to the Glutton Stratum at Tornewton Cave, though the occurrence of a late Upper 
Palaeolithic industry in the cave indicates a more recent date for most of the 

The occurrence of M. nivalis among various rodent remains excavated by J. 
Simons from Cow Cave, Chudleigh, Devon (7), suggests that there may be a pre- 
Tpswichian' deposit there. 

Hutton Cave, Somerset (10), may provide a further example of a pre-Tpswichian' 
rodent locality. One of the rodents found there, cf. Allocricetus hursae, is known 
elsewhere in the British Isles only from the Glutton Stratum of Tornewton Cave. 


A primitive form of mammoth further supports a relatively early date for the fauna 
of this cave. 

(v) 'Ipswichian' cave DEPOSITS. Sites, in addition to Tornewton Cave, include 
the hippopotamus-bearing deposits of Joint Mitnor Cave, Devon (i), with Arvicola 
cantiana-terrestris transition form (determined by W. von Koenigswald) and Microtus 
agrestis ; Minchin Hole, Glamorganshire (16), with the same species ; the nearby 
Bacon Hole (16) recently excavated by C. Stringer (1975) with Arvicola sp., 
M. agrestis, Clethrionomys glareolus and M. oeconomus ; and Alveston Fissure, 
Gloucestershire (15), with M. agrestis and Clethrionomys sp. Rodent remains from 
the interglacial site of Kirkdale Cave, Yorkshire (yy), include Arvicola sp., M. 
agrestis, Clethrionomys sp., Apodemus sylvaticus and Dicrostonyx torquatus. This 
cave was excavated over a century ago. Dicrostonyx is out of place and suggests 
some mixing of material. 

The rodent fauna of this stage is typically sparse, with Arvicola cantiana-terrestris 
transition form and Microtus agrestis the most common species. 

(vi) Caves with post-Ipswichian deposits. Cave deposits dating from the 
Last Glaciation and Holocene are far more numerous than earlier deposits, since 
there has been less time for them to be destroyed by denudation. Many examples 
are known. The distribution of rodent species at some of the more important cave 
sites believed to date from this period is shown in Table 10 (pp. 70-71 ; for refer- 
ences, see Section HI). 

The interpretation of these rodent faunas must unfortunately be undertaken with 
caution since many of the deposits in which they were found had been disturbed by 
burrowing animals ; some important rodent caves were excavated during the 
nineteenth century, so that adequate stratigraphic information is lacking. A fairly 
consistent Pleistocene faunal assemblage is nevertheless apparent, with Sciurus 
vulgaris, Muscardinus avellanarius, Apodemus flavicollis and Rattus rattus probably 
not arriving until Holocene times. A series of excavations recently conducted by 
the Peakland Archaeological Society in Derbyshire and Staffordshire (Bramwell 
i960, 1964, 1970 ; Pernetta 1966) is of special interest for the light thrown on the 
rodent faunas of the end of the Pleistocene and early Holocene. The two lemmings, 
Lemmus and Dicrostonyx, appear to have been common until the end of the Pleisto- 
cene after some of the large Pleistocene mammals, such as woolly rhinoceros and 
hyaena, had disappeared ; Microtus oeconomus seems to have survived into post- 
Pleistocene times. The apparent absence of lemmings at Levaton Cave (2, a Pleisto- 
cene site with mammoth, woolly rhinoceros, red deer, reindeer and hyaena) suggests 
that, during part of the Last Glaciation (probably an interstadial), Lemmus and 
Dicrostonyx may have been absent from the fauna, at least in the south of England. 
The Ightham fissures, Kent (Abbot 1917; Newton 1894, 1899a, b), are further 
important cave sites with rodent remains of Last Glaciation age. All the fissures 
do not appear to be contemporary, however, and there may have been some mixing 
of Holocene material. 

(vii) Caves in Scotland. A series of small caves near Inchnadamph in Scotland 
(83) are of special interest as including the most northerly British fossil rodent 


locality. From one of them, Creag nan Uamh Cave, Newton (in Peach & Home 
1917) recorded Dicrostonyx torquatus, Microtus agrestis and M. ratticeps (= M. 
oeconomus). Remains of brown bear, reindeer and lynx were also found at the same 
site. The age of these remains is uncertain, but the caves are situated in wild 
mountainous country and the possibility that they are of relatively late age needs 
to be taken into consideration. Some bones of brown bear found in a nearby cave 
have recently been dated by ^*C as only about 2700 years old (Burleigh 1972). 

(viii) Caves in Ireland. Most rodent species living in Ireland today are probably 
post-Pleistocene arrivals. Only four possibly Pleistocene species need to be con- 
sidered here. Their distribution at cave sites is shown in Table 9. 

The exact stratigraphic position of these various remains is uncertain, although it 
seems unlikely that any of the Irish mammalian faunas are earlier than Last Glaci- 
ation. Mitchell (1969) considered that the mammals could not all be attributed to 
one phase, suggesting that, although the remains of giant deer and reindeer found 
beneath the peat bogs were known to be of Late Glacial age, the Castlepook Cave 
fauna dated from an (earlier) interstadial of the Last Glaciation. A recently obtained 
i*C date for part of a mammoth bone from Castlepook Cave (33 500 ± 1200 years b.p., 
University of Dublin 122) confirms Mitchell's supposition. 

Both Lemmus and Dicrostonyx are northern species which might be expected to 
have reached Ireland in advance of other rodent species. Associated species in 

Table 9 
Distribution of rodent species at Irish cave sites 

Edenvale caves, 
Co. Clare (85) 

Caves, Co. 
Sligo (84) 
































































Lemmus lemmus 



Microtus agrestis 























Castletownroche Cave include mammoth, both mammoth and spotted hyaena being 
present in Castlepook Cave. 

Although Apodemus sylvaticits has been recorded from several cave sites, Barrett- 
Hamilton & Hinton (1910-21) observed that in some instances it was most abundant 
in the upper layers, implying a possible late arrival in Ireland. Scharff et at. (1918), 
on the other hand, considered that at Castlepook Cave Apodemus was part of the 
Pleistocene fauna. 

An imperfect skull of Microtus agrestis from Kilgreany Cave presents a problem, 
since this is the only possible fossil record of Microtus from Ireland, where this genus 
does not occur even at the present day. Savage (1966) observed that the skull is 
notably fresh and unaltered and drew attention to the occasional occurrence of vole 
skulls in north-east Ireland in owl pellets dropped by passing Scottish owls. There 
are no other records of Microtus in Ireland and the chance introduction of this 
specimen by owls or human activity needs to be taken into consideration. 

D. Other Localities 

(i) Cromerian localities. Stuart (1974) recorded Mimomys cf. savini associated 
with Dicerorhinus cf. etruscus from a site at Sugworth, near Oxford (21). He con- 
sidered that the site is unlikely to be later than Cromerian. 

(ii) HoxNiAN LOCALITIES. Carreck (1959) recorded Pitymys sp. from an inter- 
glacial tufa near Hitchin, Hertfordshire (39). This locality is unlikely to be later 
than Hoxnian in age. Other Hoxnian localities are Copford, Essex (42, Brown 
1852 ; Turner 1970), with Trogontherium hoisvilletti, and the Hoxnian type site of 
Hoxne with Trogontherium hoisvilletti (Spencer 1956), Apodemus sp., Lemmus sp., 
Arvicola sp. and Microtiis sp. (personal communication from Dr R. G. Wolff). 

(iii) Tpswichian' localities. Six important East Anglian mammalian localities, 
usually regarded as of Ipswichian age (including the type Ipswichian locality, 
Bobbitshole) have produced rodent remains as shown in Table 11. 

It has been claimed by Koenigswald (pers. comm.) that although the Arvicola 
from Barrington and from the Stuart collection from Swanton Morley (two hippo- 
potamus localities) is A. cantiana-terrestris transition form, that from the Stuart 
collection from Stutton (a locality without hippopotamus) is A . cantiana, suggesting 
affinities with the Grays-Ilford-Aveley fauna of the Thames estuary. Stuart 
(1974), on the other hand, considers that the Barrington remains include both 
'cantiana' and 'terrestris' and not merely intermediate forms, suggesting that the 
above interpretation is oversimplified. Stuart {in litt.) records A. cantiana from 
Harkstead, a locality near to and probably of the same age as Stutton. 

Other Tpswichian' localities with rodent remains are Hessle (74) with Arvicola 
and Selsey (24) with Castor. 

(iv) Last Glaciation localities. The Last Glaciation brings us within the range 
of radiocarbon dating. Four sites, two of them thus dated, will be mentioned here. 

At Upton Warren, Worcestershire (20), remains of Dicrostonyx (determined by 
Carreck) were found in association with mammoth, woolly rhinoceros and reindeer 



Distribution of rodent species at Briiil 

Sciurus vulgaris* 
Spermophilus superciliosus 
Muscardinus avellanarius* 
Castor fiber* 
Apodemus sylvaticus 
A . flavicollis* 
Rattus rattus* 
Clethrionomys glareolus 
Arvicola terrestris 
Dicrostonyx torquatus 
Lemmus lemmus 
Microtus agrestisfarvalis group 
M. oeconomus 
M. gregalis 

* Probably 



















bjo o 










p: rt 












• in 




o f- 



Lst Glaciation and Holocene cave sites 
re^rdshire Derbyshire 


'a3 S 

[g o 

72 pleistocene rodents 

Table ii 

Rodent species from East Anglian 'Ipswichian' sites 












, — - 



















, , 






1— 1 










~ — ' 





























Apodemus sylvaticus 

Clethrionomys cf . glareolus x 

Arvicola cantiana x x 

Arvicola cantiana-terrestris transition form x x x 

Arvicola sp. x 

Microtus agrestis x x x x 

M. oeconomus x 

List based on specimens in the Ipswich Museum (Stutton, Stoke Tunnel Beds, Harkstead and Bobbits- 
hole), British Museum (Natural History) (Barrington), and Stuart Collection from Swanton Morley 
(Stuart 1974). 

in terrace deposits of the River Salwarpe (Coope et al. 1961). Associated insect 
remains indicate cold continental conditions. There are ^^C dates of 41 500 ± 1200 
years (GRO 595) and 41 900 + 800 years (GRO 1245). 

At Beckford, Worcestershire (19), remains which are probably Microtus arvalis 
have been determined by Dr G. B. Corbet from specimens separated by Coope 
from material collected by D. J. Briggs from deposits of the Carrant Brook, in a 
terrace equivalent to number 2 terrace of the Warwickshire Avon or the main 
terrace of the River Severn. Associated mammals include mammoth, woolly 
rhinoceros and horse. There is a cold insect fauna and ^*C dates of 27 650 ± 250 
years (Birmingham 293) and 27 300 + 500 (Birmingham 595). 

At Fisherton, Wiltshire (22), Dicrostonyx torquatus, Microtus oeconomus and 
Spermophilus superciliosus were associated with remains of large Pleistocene 
mammals. In spite of earlier suggestions Microtus nivalis is absent. This is a 
typical Last Glaciation faunal assemblage. 

At Brean Down, Somerset (9), remains of Dicrostonyx were found in association 
with arctic fox and reindeer in talus deposits regarded by Apsimon et al. (1961) as 
being of Late Glacial age (early Zone II). 

(v) HoLOCENE LOCALITIES. Two rodent species, now extinct in the British Isles, 
survived into post-Pleistocene times. These are Castor fiber and Microtus oeconomus. 
Localities for Castor fiber include the Cambridge Fens (41), Staple Howe, Yorkshire 


(75), Star Carr, Yorkshire (76), and Thatcham, Berkshire (23). Microtus oeconomus 
has been found in the Huntspill Cut (8) in the Somerset levels and on Nornour, 
Isles of Scilly (91). 

(vi) Scottish localities. Other Scottish rodent finds include Dicrostonyx from 
Corstorphine, Edinburgh (81), and Castor fiber from the Loch of Marlee, Perthshire 
(82), and Middlestots Bog, Berwickshire (80). The various ages of these remains 
have not been determined. 

III. classification and distribution of rodents in the 


In this section rodents represented in the British Pleistocene are discussed in 
systematic order. For completeness of taxonomic reference authors and dates of 
both families and genera are given, though these are generally omitted from the 
references. In the few cases where the currently used name of an extinct species 
is based on non-British material the original reference, marked **, is given in the 
synonymy. With these exceptions, only names used for fossil specimens from the 
British Isles are cited in the synonymy and each name is given only once, with 
reference to the first paper where it appeared. The type localities of extinct species 
of rodents are indicated * in the locality lists or, if non-British, in the sections on 
general distribution. 

Locality lists for each species are divided into (a) English and Welsh, (b) Scottish 
and (c) Irish sites ; they are given in ascending stratigraphic order, as precisely as 
possible on the basis of current information. Where several records are believed 
to be of similar age they are grouped together, in alphabetical order, at the appropri- 
ate position. Numbers refer to site locations in Fig. i. Published records at 
generic level only are generally not included in the locality lists. Though coverage 
is basically restricted to the Pleistocene, Holocene records of Castor fiber and Microtus 
oeconomus are also included since both these species are now extinct in the British 
Isles. Post-Pleistocene arrivals in the British Isles are briefly mentioned for the 
sake of completeness. 

Family SGIURIDAE Brandt 1855 

Genus SCIURUS Linnaeus 1758 

Sciurus whitei Hinton 1914 

Squirrel, extinct 

1914 Sciurus whitei Hinton : 193-195, fig- 10. 

Locality : *West Runton, Norfolk (65) : Hinton 1914. 

Hinton described a new species of squirrel, 5. whitei, from a unique fourth upper 
premolar (BM(NH), M 10720) from the marine 'Monkey Gravel' overlying the 
Cromerian Upper Freshwater Bed at West Runton, Norfolk. 


Fig. 5. Distribution of remains of Sciurus 
whitei Hinton in the British Isles. 

No other fossil remains of this species have been found in the British Isles, though 
a number of fir cones from Cromerian deposits of the Forest Bed of Norfolk bear 
marks suggesting that they had been gnawed by squirrels (Newton 1882a). Newton 
(1881, 1882a, 1891) described and figured a humerus from Ostend, Norfolk (57), 
which agreed closely in form with that of the living red squirrel, 5. vulgaris Linn., 
but he was not certain whether it came from the Forest Bed or from a Recent 
alluvial deposit. 

On the continent of Europe, remains of Sciurus are known from a number of 
Early and Middle Pleistocene localities in France and Hungary, though they have 
usually been left without specific determination. Janossy (1962) discovered remains 
of a squirrel approaching 5. whitei at the Middle Pleistocene locality of Tarko in 
Hungary, and described them as belonging to a new subspecies, S. whitei hungaricus 
Janossy 1962. In his opinion 5. whitei may represent an ancestral form of 5. 

Sciurus vulgaris Linnaeus 1758 
Red squirrel 

The red squirrel is a widespread Palaearctic species, still common in parts of the 
British Isles. It occurs principally in woodland, but also in scrub beyond the 
Arctic circle. Its remains have been found in Late Pleistocene sediments of Europe 
from France to the Ukraine, but are nowhere common. 

There are no indisputable records of 5. vulgaris in the Pleistocene of the British 
Isles. Its remains are known from Dowel and Langwith Caves, Derbyshire (71, 72 : 
Bramwell i960, Mullins 1913). The Dowel Cave record is from a Holocene stratum. 
The stratigraphic position of the remains from Langwith Cave was not determined, 
but the occurrence of Rattus rattus Linn., the black rat, suggests that some Holocene 
mammalian remains were present in addition to the rich Pleistocene fauna of the 


Genus SPERMOPHILUS Cuvier 1825 [= Citellus Oken i8i6]i 

It was long suspected that the ground squirrels of the British Pleistocene represent 
two species. Hinton (in Barrett-Hamilton & Hinton 1910-21 : 724) wrote : 

Dr Forsyth Major studied the material [of ground-squirrels] with great care many years ago 
and we believe that he concluded that at least two species occur in the British Pleistocene ; 
unfortunately his results were never published. The writer in turn has made some progress 
with a similar investigation, but has not been able to complete his work yet. In his view also 
there are two species at least, both extinct, one being allied to the living Citellus [ = Spermo- 
philus] erythrogenys, the other more nearly related to C. [= Spermophilus] eversmanni. 

Following studies of specimens from the British Museum (Natural History) the 
two forms have recently been distinguished by I. M. Gromov of Leningrad {in litt.) 
as Citellus [= Spermophilus] superciliosus, from the Upper Pleistocene, and C. 
[= 5.] primigenius from deposits of earlier age. 

Spermophilus (Urocitellus) primigenius Kormos 1934 
Ground squirrel, extinct 

1876 Spermophilus ; Cheadle : 70-71. 

1882b Spermophilus Altaicus ? ; Newton : 51-54, pi. 2. 

1885 Spermophilus erythrogenoides Falconer ; Lydekker : 212-213. 

1934 ** Spermophilus primigenius Kormos : 314-315, fig. 45. 

1947 Citellus erythrogenoides Falc. ; Jackson : 168. 

Localities : Mundesley, Norfolk (60) : Newton 1882b. 

Crayford and Erith, Kent (27) : Cheadle 1876, Lydekker 1885-87, Whitaker 
1889, Newton 1890a, 1894, Barrett-Hamilton & Hinton 1910-21, Kennard 1944, 
Jackson 1947. 

Distribution in the British Isles. Remains of this species of ground squirrel 
have been found abundantly in the Middle Terrace deposits of the Thames at Cray- 
ford and Erith, Kent. Cheadle (1876) attributed remains which he had found at 
Erith to spermophilus sp. Subsequent writers (Lydekker, Whitaker, Newton, 
Barrett-Hamilton & Hinton, Kennard, Jackson) regarded the form represented at 
these localities as 5. erythrogenoides. A skull from Erith was subsequently deter- 
mined as Citellus [= Spermophilus] primigenius by Gromov, who wrote as follows 
{in litt., translation) : 

Specimen M 9605, a deformed skull with a fragment of mandible, belongs to a large form of 
Citellus [Urocitellus) primigenius Korm., approaching in its dimensions the Polish C. [U.) 
polonicus Gromov, though it preserves the basic characters of the former. The skull also has 
some characters typical for C. nogaici Top. : a well-marked basin which divides the hypo- 
and entoconid parts on the grinding surface of Mg, a weakly developed posterior crest on M^, 
a weakly developed narrowing on the hypoconid of P4, and a. foramen mentale shifted upwards. 
The second of these characters, as well as the presence of tiny cusps on the bottom of the 

^ Although the name Citellus Oken 1816 has been extensively used hitherto for these ground squirrels, 
Oken (1815-16) is not consistently binominal (Ellerman & Morrison-Scott 1966 : 3) and was rejected as 
unavailable by fiat of the International Commission on Zoological Nomenclature (Opinion 417, 1956). 


external valleys of M^-Mg, occurs also in C. polonicus. Finally, peculiar to the British 
remains is the forward shifting of the crests of the masseteric plate on the mandible. It is 
probably a distinct subspecies. 

Some isolated worn teeth of Spermophilus described by Newton (1882b : 54) as 
close to S. altaicus {= S. eversmanni) were found in deposits now regarded as of early 
Anglian age at Mundesley, Norfolk. They provide the earliest record of Spermo- 
philus in the British Isles and are provisionally referred to S. [Urocitellus) primigenius. 

General distribution. 5. primigenius (type locality Kalkberg, Nagyharsany- 
berg) is known from Villafranchian as well as from later (Giinz, Giinz-Mindel) 
deposits of Hungary. It is also present in the Middle Pleistocene of the Ukraine 
and Germany (Gromov 1965). According to Chaline (1972), ground squirrels, 
which he was unable to determine specifically, were present during the Mindel and 
Riss glaciations in France. 

The living representatives of the subgenus Urocitellus - the long-tailed ground 
squirrels 5. undulatus (Pallas 1779) and 5. parryi Richardson 1827 - inhabit steppes 
as well as meadows in the tundra-zone of central and eastern Asia and North 

Spermophilus (Colobotis) superciliosus (Kaup 1839) 

Ground squirrel, extinct 

1839 ** Spermophilus superciliosus Kaup : 112. 

1866 Spermophilus erythrogenoides (Falc.) ; Dawkins & Sanford : xxxix (nomen nudum). 

1866 Spermophilus citillus Pallas ; Dawkins & Sanford : xxxix. 

1868 Spermophilus erythrogenoides ; Falconer, in Murchison : 452-454, pi. 35. 

1974 Citellus sp. ; Stuart : 246. 

Localities : Langwith Cave, Derbyshire (72) : Mullins 1913, Barrett-Hamilton 
& Hinton 1910-21. 

Mendip Hills Caves (Bleadon Cave and others) : Dawkins & Sanford 1866, 
Falconer 1868, Stevens 1869, Sanford 1870a, b, Newton 1882a, Barrett-Hamilton 
& Hinton 1910-21, BM(NH). 

Fisherton, Wiltshire (22) : Falconer 1868, Stevens 1869, Barrett-Hamilton & 
Hinton 1910-21. 

Ightham Fissures, Kent (28) : Newton 1894, 1899b, Barrett-Hamilton & Hinton 
1910-21, BM(NH). 

Picken's Hole (layer 3), Somerset (10) : Stuart (1974). 

Pin Hole Cave, Derbyshire (73) : Jackson 1947. 

Distribution in the British Isles. 5. superciliosus is a characteristic element 
of the fauna of the Last Glaciation. 

General distribution and systematic remarks. S. superciliosus was described 
by Kaup from Eppelsheim in Germany. It is an element of the Late Pleistocene 
(Wiirm) fauna, present in almost the whole of Europe, including France, Germany, 
Poland, the European and Asiatic part of the Soviet Union, Czechoslovakia and 
Hungary. Its living relatives inhabit the steppe zone of eastern Europe and Asia. 



Fig. 6. Distribution of remains of Spermo- 
philus (Urocitellus) primigenius Kormos 
(V) and 5. (Colobotis) superciliosus (Kaup) 
(•) in the British Isles. 

In Britain it has usually been recorded as 5, erythrogenoides, a name first used by 
Falconer (in Murchison i868) for remains from caves of the Mendip Hills. Hinton 
(in Barrett-Hamilton & Hinton 1910-21 : 723) stated that recent study tended to 
show that this name must be treated as a synonym of S. superciliosus. 

I. M. Gromov of Leningrad studied the specimens from Ightham Fissures, Kent, 
and made the following remarks [in Hit., translation) : 

Two mandibular rami, no. M 11 867, are typical remains of Citellus {Colobotis) superciliosus 
Kaup, broadly distributed in the northern part of continental Europe in Late Pleistocene ; 
during some part of this time it evidently also inhabited the British Isles. From the 
characteristic features of this species, as shown by the above-mentioned material, the following 
can be stated : width of P4 no more than 11 per cent greater than its length, processus 
articularis short and broad below the head, posterior incision of the mandible relatively small, 
foramen mentale situated far from the anterior border of the masseteric plates, etc. 

Family GLIRIDAE Thomas 1897 
Genus MUSCARDINUS Kaup 1829 

Hinton (in Barrett-Hamilton & Hinton 1910-21 : 351) recorded a single tooth of 
Muscardinus from the Forest Bed of Norfolk. This specimen has not been seen by 
the present writers. Muscardinus is nevertheless known from some Cromerian 
localities on the European continent so that its presence in Britain in Cromerian 
times is not improbable. 

Muscardinus avellanarius (Linnaeus 1758) 
Common or hazel dormouse 

Although remains of this species have been found at a number of cave sites - Dog 
Holes and Pin Hole Caves (79, 73) : Jackson (1934, 1947) ; Great Doward Cave (17) : 
British Museum (Natural History) - all these remains are probably of Holocene age. 


The hazel dormouse is now distributed in southern, western and central Europe, 
including southern Sweden and Britain, and in Asia Minor. On the continent of 
Europe it has been found in interglacial deposits as early as Tiglian. It was also 
present in France during the last interglacial (Chaline 1972). 

Family CASTORIDAE Gray 1821 

Genus TROGONTHERIUM Fischer 1809 

Trogontheriutn minus Newton 1890 

Giant beaver, extinct 

i8gob Trogontherium minus Newton : 447-448. 

Localities : Red Crag of Felixstowe and *Woodbridge, Suffolk (45, 47) : Newton 
1890b, 1891, 1902, Barrett-Hamilton & Hinton 1910-21, Schreuder 1929, 1951. 

According to Schreuder (1951), who made a very thorough study of the genus 
Trogontherium, this species is more primitive than the other representatives of the 
genus, T. cuvieri Fischer and T. hoisvilleti (Laugel). 

T. minus is also known from Astian (Pliocene) sediments of Perpignan, France. 
The Red Crag specimens (like many other Crag fossils) are probably derived from 
earlier deposits and cannot be regarded as part of the contemporary Lower Pleisto- 
cene fauna. 

Trogontherium boisvilletti (Laugel 1862) 

Giant beaver, extinct 

1846 Trogontherium Cuvieri ; Owen : 184-189, figs 71-73. 

1848 Diabroticus Schmerlingi ; Pomel : 167. 

1862 ** Conodontes Boisvilletti Laugel : 715-718. 

1866 Castor trogontherium Cuvier ; Dawkins & Sanford : xxxvi. 

1902 Dipoides Lydekkeri Schlosser : 117. 

1 95 1 Trogontherium boisvilletti Laugel ; Schreuder : 403. 

1956 Trogontherium sp. ; Spencer : 354. 

Localities : Sizewell, Suffolk, and Thorpe, Norfolk^ (49, 48, Norwich Crag) : 
Lydekker 1885, Schreuder 1951. 

East Runton (64, Pastonian Forest Bed) : Newton 1882a, 1892, Barrett-Hamilton 
& Hinton 1910-21, Schreuder 1951. 

Bacton, Cromer, Kessingland, Mundesley, Overstrand, Paston and West Runton 
(58, 63, 52, 60, 62, 59, 65, 'Cromerian') : Lyell 1840, Owen 1846, Dawkins & Sanford 
1866, Owen 1869, Newton 1881, 1882a, Lydekker 1885, Reid 1890, Hinton 1914, 
Schreuder 1929, 1931. 

Clacton, Copford, Hoxne and Swanscombe (43, 42, 54, 30, Hoxnian) : Brown 1852, 
Newton 1902, Schlosser 1902, Stopes 1904, Newton 1916, Schreuder 1929, 1951, 
Spencer 1956, Sutcliffe 1964, Singer et al. 1973. 

Range : T. boisvilletti was apparently present in Britain during the Tiglian, 
Cromerian and Hoxnian. Its remains have been found frequently in deposits of 

^ See also footnote on p. 96. 



Fig. 7. Distribution of remains of Trogon- 
therium hoisvilletti (Laugel) in the British 

the Norfolk Forest Bed ; only a few Hoxnian specimens are known. These are a 
solitary incisor from Ingress Vale, Swanscombe (Newton 1902, Stopes 1904, Schreuder 
1929, 1951, Sutcliffe 1964), a cheek tooth from Copford (Brown 1852, site dated as 
Hoxnian by Turner 1970, BM(NH) 27985), a femur and eight cheek teeth from 
Hoxne (Spencer 1956) ; and part of a skull from Clacton (Singer et al. 1973). 

General distribution. T. hoisvilletti is known from the early Pleistocene in 
Britain, Holland, France (type locality Saint-Prest, near Chartres) and western parts 
of Germany. Further to the east it is, according to Schreuder (1951), replaced by a 
slightly different species, T. cuvieri Fischer, first described from the border of the 
Azov Sea. T. cuvieri was present in Germany during the Holsteinian Interglacial. 
According to Lehmann (1953), T. cuvieri is a more advanced species than T. hoisvilletti 
and may be its descendant. If this is true, the British Hoxnian remains may 
represent T. cuvieri, but there are too few suitable specimens to allow a detailed 

Genus CASTOR Linnaeus 1758 

Castor fiber Linnaeus 1758 


1846 Castor europaeus ; Owen : 190-200, figs 74-75. 

1864 Castor veterior Lankester : 355-356. 

1889 Castor fiber Linn. (= europaeus Owen) ; Whitaker : 336. 

1908b Castor plicidens Maj. ; Major : 630-635, figs 132-136. 

1964 Castor sp. ; Spencer : 338. 

Localities : Sutton and Woodbridge, Suffolk (47, Red Crag) : Lankester 1864, 
Newton 1891, Barrett-Hamilton & Hinton 1910-21, Schreuder 1929. 

East Runton (64, Pastonian Forest Bed) : Major 1908b. 

Bacton, Kessingland, Mundesley, West Runton (58, 52, 60, 65, Forest Bed) : 
Dawkins & Sanford 1866, Owen 1869, Newton 1881, 1882a, 1891, Major 1908b, 



Barrett-Hamilton & Hinton 1910-21, Hinton 1914, Schreuder 1929, 1931, Friant 

Westbury-sub-Mendip Fissure (13, lower middle Pleistocene) : Bishop 1974. 

Clacton (43, Hoxnian) : Barrett-Hamilton & Hinton 1910-21, Hinton 1923b, 
Sutcliffe 1964. 

Swanscombe (Barnfield Pit), Kent (30, Hoxnian) : BM(NH). 

Grays Thurrock and Ilford, Essex (33, 35) : Lankester 1864, Dawkins & Sanford 
1866, Whitaker 1889, Hinton 1900b, Barrett-Hamilton & Hinton 1910-21, BM(NH). 

Selsey, Sussex and Upnor, Kent (24, 31, Last Interglacial) : BM(NH). 

Kent's Cavern, Devon (6) : BM(NH). 

Cambridge Fens, Cambridgeshire ; Hay Wood Rockshelter, Somerset ; Staple 
Howe, Yorkshire ; Star Carr, Yorkshire ; Thatcham, Berks (41, 10, 75, 76, 23, 
Holocene) : Montagu 1924, Brewster 1963, Fraser & King 1954, Wymer 1962. 

Loch of Marlee, Perthshire, and Middlestots Bog, Edrom Parish, Berwickshire 
(82, 80) : Barrett-Hamilton & Hinton 1910-21. 

Fig. 8. Distribution of fossil remains of 
Castor fiber Linn, in the British Isles. 

Range. Remains of C. fiber have been found in deposits in the British Isles 
ranging in age from lowest Pleistocene (Red Crag) to Holocene. The species spread 
to Scotland but apparently never reached Ireland. Barrett-Hamilton & Hinton 
(1910-21) considered that the beaver did not become extinct in Britain before the 
thirteenth century a.d. 

General distribution. Although limited to a small number of relict localities 
as a result of extermination by man, C. fiber is still widely distributed in Europe and 
Asia. Its fossil remains are known from the Pliocene and Pleistocene of Europe and 

Systematic remarks. The beaver remains from the Red Crag were described 
by Lankester (1864) under the new name C. veterior. Newton (1891) considered that 
this species was conspecific with C. fiber and Schreuder (1929) proved it by a detailed 
comparison. C. I. F. Major (1908a) described some remains from the Forest Bed 
at East Runton as belonging to C. plicidens, a species described by him from Val 


d'Arno in Italy. Schreuder (1929) showed that the folding of tooth-enamel is a 
character depending upon the individual age of the animal and C. plicidens is there- 
fore a synonym of C. fiber. Later it was found that individuals of 'C. plicidens' 
sometimes appear among postglacial beavers. Viret (1954), however, studying the 
skull of a Villafranchian beaver from St Vallier in France, found some morphological 
differences from the Recent beaver and was inclined to retain the name C. plicidens 
for all the beavers from that period, not only for specimens with folded enamel. 
According to Lehmann (1957) these differences are rather of a subspecific character 
and he designated the Villafranchian beavers as 'Castor fiber plicidens Major 1875'. 
Until statistical analysis of the variability of the living beaver has been made, it 
seems preferable to use only the specific name Castor fiber Linn, for the Astian (Upper 
Pliocene) as well as for the Pleistocene beavers. 

Family MURIDAE Gray 1821 

Genus APODEMUS Kaup 1829 

Apodetnus sylvaticus (Linnaeus 1758) 

Wood mouse 

1846 Mus musculus (?) ; Owen : 209, fig. 79. 

1881 Mus sylvaticus ; Newton : 258-259. 

1910 Micromys {— Mus) sylvaticus (Linne) ; Jackson : 328. 

1910b Mus sp., allied to M. sylvaticus ; Hinton : 489-507. 

1915 Apodemus sp. ; Hinton : 580. 

19 1 5 Apodemus whitei Hinton : 580-581. 

1915 Apodemus sylvaticus 1^. ; Hinton : 581-582. 

Localities : West Runton (Upper Freshwater Bed), Norfolk (65) : Newton 1881, 
1882a, 1891, Barrett-Hamilton & Hinton 1910-21, Hinton 1915, BM(NH). 

Westbury-sub-Mendip Fissure, Somerset (13) : Bishop 1974. 

Hitchin, Hertfordshire (39) : Carreck 1959. 

Swanscombe (Ingress Vale), Kent (30) : Stopes 1904, Hinton 1910b, Barrett- 
Hamilton & Hinton 1910-21, Hinton 1915, Sutcliffe 1964, BM(NH). 

Hoxne, Suffolk (54) : Wolff {in lilt., Apodemus sp.). 

Grays Thurrock, Essex {^;^) : Hinton, Kennard & Newton 1900, Hinton 1915. 

Stutton, Suffolk (44) : Stuart 1974 {A. sylvaticus group). 

Bacon Hole, Glamorganshire (16) : det. A. J. Stuart {in litt.). 

Kirkdale Cave, Yorkshire {jy) : Owen 1846, Dawkins & Sanford 1866. 

Swanton Morley, Norfolk (66) : Stuart 1974 {A. sylvaticus group). 

Tornewton Cave, Devon (3) : Kowalski 1967, BM(NH). 

Aveline's Hole, Somerset (11) : Davies 1921, Hinton 1921, 1924, BM(NH). 

Dog Holes Cave, Lancashire (79) : Jackson 1910, BM(NH). 

Great Doward Cave, Herefordshire (17) : Bristol University Spelaeological 
Society, det. K. K. 

Gwaenysgor Cave, Flintshire (68) : Jackson 1947. 

Ightham Fissures, Kent (28) : Newton 1894, Barrett-Hamilton & Hinton 1910-21, 
Hinton 1915, Carreck 1957, BM(NH). 


Langwith Cave, Derbyshire (72) : Mullins 1913. 

Levaton Cave, Devon (2) : Carreck 1957. 

Pin Hole Cave, Derbyshire (73) : Jackson 1934, 1947. 

Rowberrow Cavern, Somerset (11) : BM(NH). 

Dowel Cave, Derbyshire (71) : Bramwell i960. 

Happaway Cave, Devon (5) : BM(NH). 

Lynx Cave, Denbighshire (67) : Blore 1966. 

Nazeing, Essex (37) : Hinton 1952, BM(NH). 

Joint Mitnor Cave, Devon (i, Layer 10, Holocene) : BM(NH). 

Ballynamintra Cave, Co. Waterford (90) : Barrett-Hamilton & Hinton 1910-21, 
Jackson 1929b. 

Castlepook Cave, Co. Cork (87) : Scharff, Seymour & Newton 1918, BM(NH). 

Edenvale Caves (Alice and Gwendoline, Catacombs, Newhall, Barntick), Co. Clare 
(85) : Barrett-Hamilton & Hinton 1910-21, Scharff 1906. 

Keshcorran Caves (Coffey, Plunkett), Co. Sligo (84) : Barrett-Hamilton & Hinton 
1910-21, Jackson 1929b, Scharff et al. 1903. 

Kilgreany Cave, Co. Waterford (89) : Jackson 1929b. 

Fig. 9. Distribution of fossil remains of 
Apodemus sylvaticus (Linn.) in the British 

Distribution in the British Isles. A. sylvaticus was present in the British 
Isles during the Cromerian, Hoxnian and Ipswichian interglacials. It probably 
disappeared from the country at the time of the cold stage represented by the 
Tornewton Cave Glutton Stratum. A solitary specimen from this deposit may have 
been introduced from a higher, interglacial, level. This species was present during 
the Last Glaciation and during the Holocene. Most of the localities, especially 
caves, cannot be accurately dated, and it is difficult to decide if A. sylvaticus was 
present throughout the Holocene without interruption. It may have reached Ireland 
during Upper Pleistocene times. 

General distribution. A. sylvaticus is now widely distributed in Europe, 
north Africa and western Asia. The fossil remains of this or nearly related species 
are known from the Pleistocene of Europe and China. The oldest of them are dated 


as Tiglian, with further records from deposits of Cromerian, Holsteinian and later 

Systematic remarks. Hinton (1915 : 580) referred the teeth from the Cromerian 
Upper Freshwater deposits of West Runton to Apodemus sp. Although he found 
that their pattern was indistinguishable from that of A. sylvaticus he considered 
that the material was insufficient for fine determination. From Swanscombe 
(Ingress Vale) [loc. cit.) he described a new species, which he named A. whitei, from 
part of a maxilla differing slightly from recent specimens of ^. sylvaticus. However, 
the characters concerned lie within the range of individual variability and there does 
not seem to be any good reason for regarding the West Runton form as a distinct 

Apodemus flavicollis (Melchior 1834) 
Yellow-necked mouse 

1894 Mus Ahhotti Newton : 195, pi. xi, fig. 8 (not M. abhotti Waterhouse). 

1899a Mus Lewisi Newton : 381. 

1915 Apodemus lewisi Newton ; Hinton : 582-584. 

192 1 Apodemus Flavicollis Melchior ; Hinton : 78. 

Localities : Aveline's Hole, Somerset (11) : Davies 1921, Hinton 1921, 1924. 

Dowel Cave, Derbyshire (71) : Bramwell i960. 

Etches' Cave, Derbyshire (71) : Pernetta 1966. 

Happaway Cave, Devon (5) : Hinton 1915, BM(NH). 

Ightham Fissures, Kent (28) : Newton 1894, 1899a, b, Hinton 1915, BM(NH). 

Kent's Cavern, Devon (6) : Hinton 1915. 

Pin Hole Cave, Derbyshire (73) : Jackson 1934, 1947. 

Distribution in the British Isles. Remains of A. flavicollis have been found 
almost exclusively in very late sediments. Many of the records must nevertheless 
be considered doubtful in view of the great difficulty in distinguishing remains of 
this species from those of A. sylvaticus. The remains recorded from Dowel Cave 
were found in a Neolithic layer (Bramwell i960 : [io]-pages unnumbered) and those 
from Etches' Cave were in early postglacial layers (Pernetta 1966 : 12, 15). Hinton 
(1921) considered that the Aveline's Hole remains were probably a late introduction. 
The same may be true for the Ightham Fissures. The Kent's Cavern and Happaway 
specimens were excavated a century ago. From Pin Hole Cave, Derbyshire 
(Jackson 1934), remains oi A . flavicollis have been recorded from both the superficial 
levels and at a depth of more than 3 m (10 ft), where they were apparently associated 
with Upper Pleistocene faunal remains. 

General distribution. At the present time A. flavicollis is widespread in 
Europe and east as far as the Urals and Caucasus. It is absent from a large part of 
western Europe (France, Belgium, Holland), but is present in England and southern 
Sweden. Nothing is known about the fossil remains. It is absent from the 
Pleistocene of France (Chaline 1972). It is difficult to decide whether this species 
represents a natural migration at the end of Pleistocene or during the Holocene or 
whether it was brought by man. 


Systematic remarks. According to Hinton (1915), A. lewisi, described from the 
Ightham Fissures by Newton (1899a, b), is closely related to, if not identical with, 
A. flavicollis Melchior. Jackson (1934) determined his material as 'A. fiavicollis 
{= A. lewisi)' . The lack of an anterior accessory cusp on M^ is, according to Hinton 
(1915), of no significance in specific determination, because the cusp is apparently 
absent in slightly worn teeth of A . flavicollis . It seems probable that the holotype 
of A. lewisi is nothing but a Recent specimen of A . flavicollis . 

Genus MICROMYS Dehm 1841 

Micromys minutus (Pallas 1771) 

Harvest mouse 

This species was probably introduced into Britain by man in postglacial times. 
According to Barrett-Hamilton & Hinton (1910-21) it is unknown there in a fossil 
state. Remains found at Tornewton Cave (Kowalski 1967) are from a surface 
fissure unconnected with the main sequence of deposits and are unlikely to be of any 

M. minutus, now widely distributed in Europe and Asia, is present in England 
and perhaps in the southern parts of Scotland. The genus Micromys, and probably 
the Recent species, is known in continental Europe from the late Pleistocene. 

Genus MUS Linnaeus 1758 

Mus musculus Linnaeus 1758 

House mouse 

The house mouse was introduced into Britain by man. Barrett-Hamilton & 
Hinton (1910-21) discussed some early data about its fossil occurrence in Britain 
and considered that all are erroneous or refer to Recent remains. There is no 
evidence for the occurrence of the house mouse in Britain before the Iron Age. 

Genus RATTUS Fischer 1803 

Rattus rattus (Linnaeus 1758) 

Black rat 

Although remains of the black rat have been recorded from Aveline's Hole 
(Hinton 1921) and Langwith Cave (MuUins 1913) these are probably Recent. The 
black rat was introduced into the British Isles by man during the historical period. 

Rattus norvegicus (Berkenhout 1769) 
Brown rat 

The brown rat is a recent addition to the British mammalian fauna. Jackson 
(1929b) records its remains from Kilgreany Cave, Ireland, in an upper layer with 
remains of domestic animals and prehistoric material, but these are probably of very 
recent date. 



Family CRICETIDAE Rochebrune 1883 

Genus CRICETUS Leske 1779 

Cricetus cricetus (Linnaeus 1758) 

Common hamster 

Cricetus vulgaris Runtonensis Newton : no- 113, fig. (unnumbered). 
Cricetus cricetus (Linnaeus 1758) ; Kowalski : 119- 120. 


Localities : West Runton (Upper Freshwater Bed), Norfolk (65) : Newton 1909, 
Hinton 1910b, Barrett-Hamilton & Hinton 1910-21, Osborn 1922, BM(NH). 
Tornewton Cave (Glutton Stratum), Devon (3) : Kowalski 1967, BM(NH). 

Fig. 10. Distribution of fossil remains of 
Cricetus cricetus (Linn.) in the British Isles. 

Distribution in the British Isles. Representatives of the genus Cricetus 
appeared twice in Britain : in the Cromerian interglacial and during the cold 
stage represented by the Tornewton Cave Glutton Stratum. The Tornewton Cave 
remains represent at least two individuals. 

General distribution. C. cricetus, the only living member of its genus, occurs 
at the present time in eastern Europe and in the western part of Asia. It is also 
present in western and central Europe where its range is insular and seems to be 
characterized by gradually increasing expansion. It occurs in steppes, parklands 
and meadows. In Europe its occurrence is now associated with arable fields. 
Fossil remains of Cricetus are known from the early, middle and late Pleistocene of 
western, central and eastern Europe, especially in deposits indicating a cold and arid 

Systematic remarks. All the fossil remains of the genus Cricetus are very 
uniform in their dental morphology and do not differ from the Recent ones. They 
do, however, vary greatly in size. On this basis many new forms have been described, 
some under specific, others under subspecific names. The specimen from the 
Cromer Forest Bed, BM(NH) M18352, an upper jaw with a tooth-row 9-3 mm long. 



is the holotype of a form known as Cricetus runtonensis or C. cricetus runtonensis. 
According to Schaub (1930), it is a synonym of C. c. major Woldfich described from 
the late Pleistocene deposits of Czechoslovakia. Other palaeontologists use the 
name C. runtonensis to designate larger representatives of Cricetus from the early 
Pleistocene, usually without giving sufficient morphological reasons for its distinctness 
from later populations. According to Fahlbusch (1969), there are some peculiarities 
in the pattern of Mg in C. runtonensis, but a similar morphology can also be found 
in some Recent specimens. In the opinion of one of us (K. K.) the large early 
Pleistocene hamsters are at most subspecifically distinct from those of the late 
Pleistocene and present day. The great variability of this species in the Pleistocene 
was connected with changing climatic conditions. 

Genus ALLOCRICETUS Schaub 1930 

cf. Allocricetus bursae Schaub 1930 

Hamster, extinct 

1870a Cricetus songarus ; Sanford : 51, 56, fig. 6. 

1870b Cricetus {Mus) songarus (Pallas) ; Sanford : 128-129 '< pl- viii, fig. 6. 

1913 Phodopus sanfordi Hinton, in Barrett-Hamilton & Hinton 1910-21 : 383, fig. 53. 

1967 Phodopus songorus (Pallas) ; Kowalski : 113-114. 

Localities : Hutton Cave, Somerset (10) : Sanford 1870a, b, Newton 1909, 
Barrett-Hamilton & Hinton 1910-21, BM(NH). 

Tornewton Cave (Glutton Stratum), Devon (3) : Kowalski 1967, BM(NH). 

Fig. II. Distribution of remains of cf. 
Allocricetus bursae Schaub in the British 

Distribution in the British Isles. The occurrence of this species in the 
Glutton Stratum of Tornewton Cave shows that it was present during the pre- 
hippopotamus cold stage represented by this deposit. Although the age of the 
Hutton Cave fauna is unknown, the occurrence of remains of a relatively primitive 


form of mammoth suggests that this record is at least as early as that from Tornewton 

General distribution and systematic remarks. Small representatives of 
Cricetidae are common in the early, middle and late Pleistocene of Europe. The 
uniform structure of the teeth in Cricetidae makes the systematic determination of 
their remains very difficult. According to Schaub (1930), the small hamsters of the 
late Pleistocene of Europe should be referred to Phodopus songorus. Janossy (1961) 
later stated that Schaub based his determination on incorrectly determined Recent 
specimens and claimed that the small hamster described by him is conspecific with 
Recent Cricetulus migratorius. The same name was used for English specimens by 
Kurten (1969). Chaline (1972) did not find this species in France, but determined 
all late Pleistocene small French hamsters as Allocricetus bursae Schaub 1930. A 
more detailed study of English specimens will be necessary before it is possible to 
decide whether they belong to one or two species and what species there are. 

In the earliest description of British hamster remains (those from Hutton), 
Sanford (1870) applied the name Cricetus songarus. Hinton (in Barrett-Hamilton & 
Hinton 1910-21 : 383) considered that the remains should be placed in the genus 
Phodopus and that they could not be synonymized with P. songorus but needed a 
new name, for which he proposed P. sanfordi. No reasons for this decision are 
given. If the British specimens are conspecific with A. bursae Schaub then this 
name is a junior synonym of P. sanfordi Hinton and must be replaced by it. 

Small hamsters are now an element of the steppe fauna, and their species are 
broadly distributed in south-eastern Europe as well as in western and central Asia. 
C. migratorius ranges from south-eastern Europe to China. This species, partly 
cited under the name of A. bursae Schaub, is known as a fossil from the late Pleisto- 
cene of Germany, Switzerland, Spain, France, Czechoslovakia, Hungary, Poland 
and the Soviet Union. 

Genus DICROSTONYX Gloger 1841 

Dicrostonyx torquatus (Pallas 1779) 

Collared lemming 

1869 Lemmus torquatus ; Stevens : no, 113. 

1870a Arvicola Gulielmi Sanford : 51, 55, fig. 2. 

1870b Arvicola Gulielmi Sanford : 125, pi. viii, fig. 2. 

1874 Myodes torquatus Pall. ; Blackmore & Alston : 469-470. 

1910a Dicrostonyx gulielmi Sanford ; Hinton : 38-39. 

1910a Dicrostonyx henseli Hinton : 37-38. 

i960 Dicrostonyx sp. ; Bramwell : [10]. 

Localities : Westbury-sub-Mendip, Somerset (13) {Dicrostonyx s^., lower middle 
Pleistocene) : Bishop 1974. 

Erith, Kent (27) : Newton 1890a, Hinton 1910b, Barrett-Hamilton & Hinton 
1910-21, Hinton 1926b, Kennard 1944, Jackson 1947. 

Hutton Cave, Somerset (10) : Stevens 1869, Sanford 1870a, b, Blackmore & Alston 
1874, Jackson 1909, Barrett-Hamilton & Hinton 1910-21, Hinton 1926b, BM(NH). 


Tornewton Cave (Glutton Stratum and Reindeer Stratum), Devon (3) : Kowalski 
1967, BM(NH). 

Aveline's Hole, Somerset (11) : Davies 1921, Hinton 1921, 1924, 1926b, BM(NH). 

Chudleigh Fissure, Devon (7) : Hinton 1926b. 

Dog Holes Cave, Lancashire (79) : Jackson 1909, 1910, 1929a, Barrett-Hamilton & 
Hinton 1910-21, Hinton 1926b, Jackson 1947, BM(NH). 

Elder Bush Cave, Staffordshire (69) : Bramwell 1964. 

Fisherton, Wiltshire (22) : Stevens 1869, Blackmore & Alston 1874, Jackson 1909, 
Barrett-Hamilton & Hinton 1910-21, Hinton 1926b, BM(NH). 

Great Doward Cave, Herefordshire (17) : Bristol University Spelaeological 
Society, det. K. K. 

Gwaenysgor Cave, Flintshire (68) : Jackson 1932 {fide Jackson 1947), Jackson 1947. 

Ightham Fissures, Kent (28) : Newton 1894, Jackson 1909, Hinton 1910a, b, 
Barrett-Hamilton & Hinton 1910-21, Hinton 1926b, Jackson 1929a, 1947, Zimmer- 
mann 1959, BM(NH). 

Kent's Cavern, Devon (6) : BM(NH). 

King Arthur's Cave, Herefordshire (17) : BM(NH) (Hinton collection). 

Langwith Cave, Derbyshire (72) : Mulhns 1913, Hinton 1910a, Barrett-Hamilton 
& Hinton 1910-21, Hinton 1926b, Jackson 1929a, 1947, BM(NH). 

Merlin's Cave, Herefordshire (18) : Hinton 1925, 1926b, BM(NH). 

Murston, Kent (32) : Newton 1890a, Barrett-Hamilton & Hinton 1910-21. 

Pin Hole Cave, Derbyshire (73) : Jackson 1934, 1947. 

Rowberrow Cavern, Somerset (11) : BM(NH) (Hinton collection). 

Upton Warren, Worcestershire (20) : Coope et al. 1961. 

Brean Down, Somerset (9) : Apsimon et al. 1961. 

Fox Hole Cave, Derbyshire (71) : Bramwell 1970. 

Dowel Cave, Derbyshire (71) : Bramwell i960. 

Lynx Cave, Denbighshire (67) : Blore 1966. 

Etches' Cave, Derbyshire (71) : Pernetta 1966. 

Hackney, London (36) : BM(NH). 

Angel Road, Middlesex (36) : Hinton 1912, Barrett-Hamilton & Hinton 1910-21, 
Hinton 1926b. BM(NH). 

Nazeing, Essex (37) : Hinton 1952, BM(NH). 

Ponders End, Middlesex (36) : Hinton 1926b. 

Corstorphine, nr. Edinburgh, Scotland (81) : Evans 1907, 1913, Barrett-Hamilton 
& Hinton 1910-21, Jackson 1929b. 

Creag nan Uamh Cave, Sutherland (83) : Peach & Home 1917. 

Ballynamintra Cave, Co. Waterford, Ireland (90) : Coleman 1965. 

Castlepook Cave, Co. Cork {Sj) : Ussher et al. 1908, Jackson 1910, Ussher 1910, 
Hinton 1910a, Barrett-Hamilton & Hinton 1910-21, Hinton 1926b, Jackson 1929b, 
1947, Coleman 1965. 

Castletownroche Cave, Co. Cork (88) : Coleman 1965. 

Edenvale Caves (Alice and Gwendoline, Catacombs, Newhall), Co. Clare (85) : 
Scharff 1906, Hinton 1926b, Coleman 1965, BM(NH). 


Keshcorran Caves (Coffey, Plunkett), Co. Sligo (84) : Jackson 1909, Scharff et al. 
1903, Scharff, Seymour & Newton 1918, Barrett-Hamilton & Hinton 1910-21, 
Hinton 1926b, Jackson 1929b, Coleman 1965, BM(NH). 

Kilgreany Cave, Co. Waterford (89) : Jackson 1929b, Coleman 1965. 

Red Cellar Cave, Co. Limerick (86) : Coleman 1965. 

Brandon, Warwickshire, and Penkridge, Staffordshire, are two additional Dicro- 
stonyx localities not marked on Fig. i, which were notified to us by Mr J. Carreck 
[in Hit.) after the compilation of the above list. 

Fig. 12. Distribution of fossil remains of 
Dicrostonyx tovquatus (Pallas) in the 
British Isles. 

Distribution in the British Isles. Dicrostonyx is known in the British Isles 
from at least three stages of the Pleistocene. The earliest record [Dicrostonyx sp.) 
is from the lower middle Pleistocene site of Westbury-sub-Mendip. The second 
stage is represented by remains from the Corbicula bed at Erith and the Glutton 
Stratum of Tornewton Cave. At the last-mentioned site Dicrostonyx disappeared 
in the overlying Last Interglacial deposits and reappeared again in the Last Glaci- 
ation Reindeer Stratum. During the Last Glaciation it reached its greatest climax, 
spreading throughout England, Wales, Scotland and Ireland. Its remains are 
especially abundant in deposits dating from the later part of this stage and it was 
clearly a dominant element of the tundra faunal assemblage of that time. At 
Nazeing, Essex, it apparently survived until late Glacial pollen Zone III. 

General distribution. The collared lemming is now widely distributed in the 
tundra belt of the Holarctic, in both Eurasia and North America. The American 
subspecies are sometimes recognized as specifically different from those of Eurasia. 
Fossil remains are known from the deposits of four different glaciations in Europe 
and they are also present in Asia and North America. Pre-Cromerian collared 
lemmings from Czechoslovakia and Poland are morphologically different from later 
ones and were named Dicrostonyx simplicior Fejfar 1966. During the Last Glaciation 
D. torquatus in Europe reached as far south as Hungary, Switzerland and central 


Systematic remarks. Hinton (1910a) distinguished two species of fossil 
collared lemmings in the British Isles, Dicrostonyx gulielmi and D. henseli. Accord- 
ing to him the differences between these were as great as those between different 
Recent species of the genus Dicrostonyx. It is now generally accepted, however, 
that disregarding the Canadian D. hudsonius (which does not concern us here) all 
living forms belong to one species. The European fossil forms may, at the most, 
be subspecifically different from Recent ones, but this seems not to be the case as the 
two forms described by Hinton are present together at nearly all localities. Janossy 
(1954) found in Hungarian caves not only all possible intermediate forms between 
D. gulielmi and D. henseli, but also specimens with left tooth-row of one type and 
right one of the other. Most palaeontologists are of the opinion that there was only 
one species of the collared lemming in the Upper Pleistocene of Europe and that it 
was conspecific with the living D. torquatus. Like the Recent population, it was 
very variable (Mandach 1938). Recently Agadzhanian (1973) has stated that, 
among the populations of Recent and fossil collared lemmings, there are different 
morphotypes, differing in the grade of complication of the crown pattern of their 
molars. He considered that the simpler morphotypes are more numerous among 
the populations from the Last Glaciation of Eurasia than among Recent animals from 
north Asia. For this reason he favoured preserving the name D. gulielmi as the 
oldest available designation for the collared lemmings from the time of the Last 
Glaciation of Eurasia. 

Genus LEMMUS Link 1795 

Lemmus lemmus (Linnaeus 1758) 

Norwegian lemming 

1870a Lemmus norvegicus (var.) ; Sanford : 51, 56, fig. 4. 

1870b Lemmus norvegicus ; Sanford : 125-126, pi. viii, fig. 4. 

1874 Myodes lemmus (Linn.) ; Blackmore & Alston : 470-471. 

1921 Lemmus lemmus Linn. ; Hinton : 75-76. 

1950 Lemmus sp. ; Schreuder : 629, 633-634. 

Localities : Westbury-sub-Mendip, Somerset (13) {Lemmus sp., lower middle 
Pleistocene) : Bishop 1974. 

Hoxne, Suffolk (54) : Wolff [in litt.). 

Swanscombe (Barnfield Pit, Upper Middle Gravel), Kent (30) {Lemmus sp.) : 
Schreuder 1950. 

Erith, Kent (27) : Newton 1890a, Hinton 1910b, Barrett-Hamilton & Hinton 
1910-21, Hinton 1926b, Kennard 1944, Jackson 1947, BM(NH). 

Hutton Cave, Somerset (10) : Sanford 1870a, b, Blackmore & Alston 1874, 
Jackson 1909, Barrett-Hamilton & Hinton 1910-21, Hinton 1926b, BM(NH). 

Tornewton Cave (Glutton Stratum), Devon (3) : Kowalski 1967, BM(NH). 

Aveline's Hole, Somerset (11) : Davies 1921, Hinton 1921, 1926b, BM(NH). 

Chudleigh Fissure, Devon (7) : Hinton 1926b. 

Dog Holes Cave, Lancashire (79) : Jackson 1909, 1910, Barrett-Hamilton & 
Hinton 1910-21, Hinton 1926b, BM(NH). 


Elder Bush Cave, Staffordshire (69) : Bramwell 1964. 

Great Doward Cave, Herefordshire (17) : Bristol University Spelaeological 
Society, det. K. K. 

Ightham Fissures, Kent (28) : Newton 1894, Bate 1901, Jackson 1909, Hinton 1910a, 
Barrett-Hamilton & Hinton 1910-21, Hinton 1926b, Zimmermann 1959, BM(NH). 

Kent's Cavern, Devon (6) : Kennard 1945-46. 

Langwith Cave, Derbyshire (72) : Mullins 1913, Barrett-Hamilton & Hinton 
1910-21, Hinton 1926b. 

Merlin's Cave, Herefordshire (18) : Hinton 1924, 1926b, BM(NH). 

Pin Hole Cave, Derbyshire (73) : Jackson 1934, 1947. 

Uphill Cave, Somerset (9) : Barrett-Hamilton & Hinton 1910-21, Hinton 1926b, 

Dowel Cave, Derbyshire (71) : Bramwell i960. 

Etches' Cave, Derbyshire (71) : Pernetta 1966. 

Harborough Cave, Derbyshire (70) : Jackson 1929a. 

Nazeing, Essex (37) : Hinton 1952, BM(NH). 

Castlepook Cave, Co. Cork, Ireland {8y) : Ussher et al. 1908, Ussher 1910, Scharff, 
Seymour & Newton 1918, Barrett-Hamilton & Hinton 1910-21, Jackson 1929b, 
Coleman 1965, BM(NH). 

Castletownroche Cave, Co. Cork (88) : Coleman 1965. 

After the compilation of the above list we have been informed by Mr J. Carreck 
{in litt.) that he has also determined remains of Lemmus lemmus from deposits of 
Ipswichian (Zone Hb) age at Wretton, Norfolk (not marked on Fig. i). 

Fig. 13. Distribution of fossil remains of 
Lemmus lemmus (Linn.) in the British Isles. 

Distribution in the British Isles. L. lemmus is distributed in Pleistocene 
sediments of England and southern Ireland. In many localities it has been found 
together with Dicrostonyx torquatus, but it is usually less numerous. Most of the 
fossil localities with Lemmus are of Last Glaciation age. Its occurrence at Nazeing, 
where it was found in peaty muds lying deeper than the calcareous muds in which 


Dicrostonyx torquatns appears for the last time, seems to be the latest well-dated 
evidence of its presence in Britain. This may indicate that Lemmus disappeared 
slightly earlier than Dicrostonyx, though it could also be explained by lack of 
sufficient specimens. 

Lemmus is known from at least four stages of the British Pleistocene. Earlier 
records are the lower middle Pleistocene site of Westbury-sub-Mendip, the Hoxnian 
of Hoxne and Swanscombe (Barnfield Pit, Upper Middle Gravel), and the stage or 
stages represented by the Corbicula Bed of Crayford and Erith and the Glutton 
Stratum of Tornewton Cave. 

General distribution. The genus Lemmus has a circumpolar distribution 
ranging in the Old World from Scandinavia to the Bering Strait and including the 
arctic regions of North America. The differences between the Eurasiatic and 
American forms do not justify their distinction as different species (Sidorowicz 
1964) : they can all be treated as subspecies of a single species, L. lemmiis. Lemmus 
coexists in a great part of its range with Dicrostonyx, but is generally more southerly 
in distribution : it does not occur in Greenland or on the Canadian islands north of 
Viscount Melville Sound, where Dicrostonyx is abundant. 

The genus Lemmus appeared very early in Europe. Its remains are known from 
middle Villafranchian deposits of Hungary, Poland and Germany. During the 
early Pleistocene it extended further south in eastern Europe than Dicrostonyx, 
and was present, for example, in Romania, where Dicrostonyx never penetrated. Its 
remains have been found at most early and middle Pleistocene fossil localities of 
central and western Europe and the genus is a characteristic element of the arctic 
fauna of the last two glaciations. 

Systematic remarks. Fossil remains of Lemmus from the late Pleistocene of the 
British Isles and from the continent of Europe are indistinguishable, on the basis of 
teeth and preserved skeletal remains, from those of Recent L. lemmus. Villa- 
franchian to middle Pleistocene remains are specifically different from the Recent 

Genus CLETHRIONOMYS Tilesius 1850 

Clethrionomys glareolus (Schreber 1780) 

Bank vole 

1846 Arvicola pratensis ; Owen : 208, fig. 78. 

1870a Arvicola glareolus (= pratensis) ; Sanford : 56. 

1870b Arvicola glareolus (Schreber) = pratensis (Baillon) = riparia (Yarrell) ; Sanford : 124. 

1882a Arvicola [Evotomys Coues & Allen) glareolus Schreber ; Newton : 82-83, P^- xiv, 
fig. i-ic. 

1900 Microtus {Evotomys) glareolus Schreb. ; Hinton, Kennard & Newton : 347-349. 

1 90 1 Microtus glareolus ; Hinton : 142. 
1910b Evotomys sp. ; Hinton : 492, 494, 497. 

1926 Evotomys harrisoni Hinton : 216-217, pi. vii, fig. 2. 

1926 Evotomys kennardi Hinton : 225-226, pi. vii, fig. 3. 

1950 Clethrionomys sp. ; Schreuder : 629, 634-635. 


Localities : West Runton (Upper Freshwater Bed), Norfolk (65) : Blackmore & 
Alston 1874, Newton 1882a, 1891, Hinton 1910b, Osborn 1922, Hinton 1926b, 

Westbury-sub-Mendip, Somerset (13) : Bishop 1974. 

Clacton, Essex (43) : Singer et al. 1973. 

Hitchin, Hertfordshire (39) : Carreck 1959. 

Swanscombe (Barnfield Pit, Upper Middle Gravels and Ingress Vale), Kent (30) : 
Stopes 1904, Hinton 1926b, Schreuder 1950, Sutcliffe 1964, Carreck {in litt.). 

Aveley, Essex (34) : BM(NH). 

Grays Thurrock, Essex (33) : Hinton, Kennard & Newton 1900, Hinton 1901, 
1910b, 1926b, BM(NH). 

Northfieet, Kent (29) : det. A. J. Stuart {in litt.). 

Hutton Cave, Somerset (10) : Sanford 1870a, b, Blackmore & Alston 1874. 

Tornewton Cave (Glutton Stratum and Reindeer Stratum), Devon (3) : Kowalski 
1967, BM(NH). 

Alveston Fissure, Gloucestershire (15) : det. G. B. Corbet. 

Bacon Hole, Glamorganshire (16) : Stuart {in litt.). 

Kirkdale Cave, Yorkshire {yj) : BM(NH). 

Swanton Morley, Norfolk (66) : Stuart 1974. 

West Wittering, Sussex (24) : BM(NH). 

Aveline's Hole, Somerset (11) : Davies 1921, Hinton 1921, 1924, BM(NH). 

Brixham Cave, Devon (4) : Lydekker 1885, Hinton 1926b, BM(NH). 

Chudleigh Fissure, Devon (7) : BM(NH). 

Dog Holes Cave, Lancashire (79) : Jackson 1910, BM(NH). 

Elder Bush Cave, Staffordshire (69) : Bramwell 1964. 

Great Doward Cave, Herefordshire (17) : BM(NH). 

Gwaenysgor Cave, Flintshire (68) : Jackson 1947. 

Ightham Fissures, Kent (28) : Newton 1894, Hinton 1910b, 1926b, Zimmermann 
1959, BM(NH). 

Kent's Cavern, Devon (6) : Owen 1846, Dawkins & Sanford 1866, Blackmore & 
Alston 1874, Lydekker 1885, Kennard 1945-46, BM(NH). 

Langwith Cave, Derbyshire (72) : MuUins 1913. 

Merlin's Cave, Herefordshire (18) : Hinton 1925. 

Pin Hole Cave, Derbyshire (73) : Jackson 1934, 1947. 

Rowberrow Cavern, Somerset (11) : BM(NH). 

Dowel Cave, Derbyshire (71) : Bramwell i960. 

Happaway Cave, Devon (5) : BM(NH). 

Nazeing, Essex (37) : Hinton 1952, BM(NH). 

Joint Mitnor Cave (layer X, Holocene), Devon (i) : BM(NH). 

Distribution in the British Isles. The bank vole appeared in England for 
the first time during the Cromerian Interglacial. It was also present during the 
Hoxnian (Hitchin, Swanscombe). Although a few remains were found in the pre- 
hippopotamus Glutton Stratum of Tornewton Cave, this species is lacking from 
typical faunal assemblages of this time, for example from Clevedon Cave. It is a 
common element in deposits of Last Interglacial, Last Glaciation and Holocene age. 



Fig. 14. Distribution of fossil remains of 
Clethrionomys glareolus (Schreber) in the 
British Isles. 

Clethrionomys glareolus is an element of the forest fauna. It was probably not 
present in this country during the maxima of the giaciations, though it was able 
to reach it very early after the retreat of the ice and invaded the British Isles many 
times in all the interglacials and in post-glacial time. 

General distribution. C. glareolus is now distributed throughout Europe, 
northern and central Asia and probably North America. Remains of the genus 
Clethrionomys are common in faunas of early middle Pleistocene age from continental 
Europe and have been described under different specific names, though they probably 
all belong to C. glareolus. C. glareolus is abundant in the interglacial and inter- 
stadial deposits of the late Pleistocene of Europe. 

Systematic remarks. Only a few remains of Clethrionomys are known from the 
Norfolk Forest Bed. Although they were determined by Hinton (1926b) as 'Evo- 
tomys sp.', they cannot be distinguished from Recent C. glareolus. The bank vole 
is a primitive, and very conservative, species of vole and its existence in Europe in 
the early Pleistocene is not unexpected. 

Hinton (1926b) described from the Ightham Fissures two new species of Clethrio- 
nomys, 'Evotomys kennardi' and 'E. harrisoni', both from the same faunal assemblage. 
The material from Ightham is however a mixture of late Glacial and Recent bones 
and the holotype of E. harrisoni is a well-preserved skull of a typical C. glareolus. 
It is the skull of a young individual and its supposed characters (small dimensions, 
broad inter-orbital region) are related to its juvenile condition ; it is probably a 
Recent specimen. 'E. kennardi' seems to be of earlier geological age and its dimen- 
sions are somewhat larger than those of Recent specimens. According to Hinton 
(1926b), it belongs to the 'E. nageri' group. C. g. nageri is now generally recognized 
as an Alpine subspecies (or perhaps only an ecotype) of C. glareolus, living under a 
more severe climate. It seems probable that during the difficult conditions of the 
Last Glaciation individuals of Clethrionomys grew bigger than during interglacial 
and postglacial times, but there are no reasons to determine them as a new species. 




Genus PLIOMYS Mehely 1914 

Pliomys episcopalis Mehely 19 14 

Vole, extinct 

** Pliomys episcopalis Mehely : 195-203, pis 4-5. 
Pliomys episcopalis Mehely ; Bishop : 309, 314. 


Locality : Westbury-sub-Mendip Fissure, Somerset (13) : Bishop 1974, 1975. 

Fig. 15. Distribution of remains of Pliomys 
episcopalis Mehely in the British Isles. 

Distribution in the British Isles. Pliomys is known from only one British 
locality, Westbury-sub-Mendip, where it occurs in association with a lower middle 
Pleistocene fauna. 

General distribution. P. episcopalis (type locality Betfia in Romania) appeared 
in the late Villafranchian of Europe and became widespread in the lower middle 
Pleistocene. In Italy it probably survived until the Holsteinian. Its remains are 
distributed from the European part of the Soviet Union to France and Italy. This 
species is generally recognized as an element of the steppe fauna. 

Genus MIMOMYS Major 1902 

Mimomys pliocaenicus Major 1902 

Vole, extinct 

1874 Arvicola amphibius (Linn.) ; Blackmore & Alston : 462-464 [partim). 

1882a Arvicola {Evotomys) intermedins Newton : 83, pi. 13 [partim). 
1902 Mimomys pliocaenicus Major : 102-107, figs 13-15. 

Distribution in the British Isles. Although one rolled microtine tooth from 
the Red Crag has provisionally been referred to Mimomys by Spencer (1964) no 
remains identifiable at specific level are known from Britain earlier than the Norwich 
Crag. M. pliocaenicus has been found in the Norwich Crag and Weybourne Crag 



Fig. i6. Distribution of remains of 
Mimomys pliocaenicus Major in the British 

of Covehithe, Easton Bavents and Sizewell, Suffolk, and Thorpe ^ and Bramerton, 
Norfolk (51, 50, 49, 48, 55). Its remains are also known from deposits of uncertain 
age at Kyson, Suffolk (47), and from the Weybourne Crag and probably Pastonian 
Forest Bed deposits at East Runton, Norfolk (64). All the specimens from this 
last locality seen by us in the British Museum (Natural History) are rounded, with 
visible traces of transport by water. 

Bibliography. Blackmore & Alston 1874, Newton 1882a, Major 1902, Hinton 
1910b, Barrett-Hamilton & Hinton 1910-21, Hinton 1926b, Cranbrook 1955a, b, 
Carreck 1966, West 1968. Specimens have been seen in the BM(NH) and Ipswich 

General distribution. M. pliocaenicus was first described by C. I. F. Major 
from the Val d'Arno in Italy. It is known from Late Villafranchian (Tiglian) 
deposits in France, Holland, Belgium, Germany, Poland, Hungary, Czechoslovakia, 
Romania and the Soviet Union (including Siberia). It has not been found in 
Cromerian (s.l.) faunal assemblages outside Britain, where it is of Pastonian age. 

Systematic remarks. M. pliocaenicus is a characteristic species. It developed 
from smaller and more primitive ancestors living in the early Villafranchian [M. 
polonicus Kowalski) and disappeared or rather evolved into other species at the end 
of the Tiglian Interglacial. 

^ There are several places in East Anglia known as Thorpe and the location of the M. pliocaenicus 
site is not proved beyond doubt. The two most likely localities are Thorpe near Norwich, Norfolk, and 
Thorpe near Aldeburgh, Suffolk. C. I. F. Major (1902) described specimens collected by Mr Fitch from 
Thorpe, preserved in the Norwich Museum, though he did not indicate the county in which the Thorpe 
in question is situated. Carreck (1966) assumed that it was the Thorpe near Aldeburgh. Mr Fitch is 
known to have collected in the Norwich Crag of the Norwich area, however, and as the specimens sub- 
sequently found their way to Norwich Museum the possibility that they came from the Thorpe near 
Norwich must also be considered. Mr H. E. P. Spencer {in lift.) favours the last-mentioned locality, 
since there is a well-documented Crag mammal site there, whereas he points out that Thorpeness in 
Suffolk is on Coralline Crag. Mr P. Cambridge informs us of a Norwich Crag pit at Shell Cottages, 
Thorpe, Aldringham, near Aldeburgh, Suffolk. He too thinks the Thorpe near Norwich, where there were 
formerly several Norwich Crag pits with mammalian remains, is the most likely locality for the remains 
of M. pliocaenicus described by Major. We have tentatively accepted the Norfolk alternative, and this 
is marked on Fig. i as locality 48. 


Mimotnys reidi Hinton 1910 
Vole, extinct 

1882a Arvicola {Evotomys) intermedius Newton : 83 (partim). 
1910b Mimomys reidi Hinton : 491. 

Distribution in the British Isles. This species is known in Britain from the 
Weybourne Crag at *Trimingham (61) and from the Norwich Crag at Sizewell (49), 
In the collections of the British Museum (Natural History) there are specimens 
from the Savin collection labelled 'West Runt on, Upper Freshwater Bed', but they 
are probably mislabelled or were found in a secondary layer. Hinton (1926b) 
stated that this species was not found outside the Norwich and Weybourne Crags. 

Bibliography. Newton 1882a, Barrett-Hamilton & Hinton 1910-21, Hinton 
1910b, 1926b. Specimens have been seen at the BM(NH) and Ipswich Museum. 

General distribution. M. reidi (probably identical with M. petenyi Mehely 
1914) is known from many localities of Tiglian age in Holland, France, Germany, 
Italy, Poland, Czechoslovakia, Hungary and the Soviet Union (including Siberia). 

Systematic remarks. The holotype of this species was described from the 
Weybourne Crag at Trimingham, Norfolk, but the systematic position of the remains 
determined as M. reidi from continental Europe presents a problem : different 
forms, probably different stages of the same phyletic line, appear to be represented. 

Fig. 17. Distribution of remains of Mzwomys 
reidi Hinton (v) and M. newtoni Major (#) 
in the British Isles. 

Mimomys newtoni Major 1902 

Vole, extinct 

1902 Mimomys newtoni Major : 102-107, figs 13-15. 

Distribution in the British Isles. In Britain M. newtoni is known from 
deposits ranging in age from Norwich Crag to either Weybourne Crag or Forest 
Bed. The holotype was described by Major (1902 : text-fig. 13, no. 7) as having 


been found in the 'East Runton Forest Bed'. It is now known that the deposits 
at East Runton are Pastonian and the holotype is most likely to be of this age. 
M. newtoni is also known from specimens in the British Museum (Natural History) 
collected by the Earl of Cranbrook from the Norwich Crag of Easton Bavents, 
Suffolk (50). The specimen in the BM(NH) from the Norwich Crag at Bramerton, 
Norfolk, (Hinton 1926b : 376, footnote) 'which may be referred to M. newtoni' is 
clearly different from the holotype and probably does not belong to the same 

Bibliography, Major 1902, Hinton 1910b, Barrett-Hamilton & Hinton 1910-21, 
Hinton 1926b. 

General distribution. M. newtoni is known outside Britain from deposits of 
Tiglian age in Holland, France, Germany, Poland, Czechoslovakia, Hungary and 
the Soviet Union. 

Systematic remarks. Small representatives of the genus Mimomys Major are 
so rare in the Lower Pleistocene strata of East Anglia that it is difficult to decide 
how many species are represented. Some of the specimens of 'M. newtoni' from 
East Runton (M6967, numbers 19, 20 and 23) are visibly larger than the holotype 
(Mj 2-9 and 3-0 mm long, compared with 2-5 mm) and have cement in the re-entrant 
angles, absent in the type specimen. There may be two 'chronospecies' of different 
geological ages. The continental specimens (e.g. from Kadzielnia, Poland) are 
identical with these larger specimens of 'M. newtoni' and different from the holotype. 
Further research will be necessary to determine whether an additional species should 
be established. 

Mimomys savini Hinton 1910 

Vole, extinct 

1874 Arvicola amphibius (Linn.) ; Blackmore & Alston : 462-464 [partim). 

i88i Arvicola {Evotomys, Coues) intermedia Newton : 258. 

1902 Mimomys intermedins (Newt.) ; Major : 102-107. 

1910b Mimomys savini Hinton : 491. 

1910b Mimomys majon Hinton : 491. 

1958 Mimomys milleri Kretzoi : 55. 

Distribution in the British Isles. M. savini is a common species in the 
Upper Freshwater Bed at West Runton (65) and it is also known from East Runton 
(64). According to Newton (1882a), it has also been found at Cromer (63), Geldeston 
(53) and Kessingland (52). Mimomys cf. savini was recorded by Stuart (1974) from 
Cromerian deposits at Sugworth, near Oxford (21). 

Bibliography. Blackmore & Alston 1874, Newton 1881, 1882a, 1891, Reid 
1890, Major 1902, Hinton 1910b, Barrett-Hamilton & Hinton 1910-21, Hinton 
1920, 1926b, Kretzoi 1958, 1965, Pasquier 1972. 

General distribution. Mimomys savini is widely distributed at localities of 
'Cromerian' age in Holland, France, Germany, Italy, Poland, Czechoslovakia, 
Yugoslavia and the Soviet Union (including Siberia). 


Fig. 18. Distribution of remains of Mimowys 
savini Hinton in the British Isles. 

Systematic remarks. M. savini was described by Newton (1881) from the 
Upper Freshwater Bed of *West Runton under the name of Arvicola intermedia. 
Hinton (1910b) described two further species of the same genus, M. majori and M. 
savini, from the same deposit. According to him, they differed from M. intermedius 
only in sUght differences in the pattern of M^, the dimensions of all the molars and 
the patterns of all other teeth being identical. Kretzoi (1958) found that A. inter- 
media is a junior homonym and must therefore be replaced by the name Mimomys 
milleri Kretzoi ; in a later paper (1965) he showed that the three species M. inter- 
medius, M. savini and M. majori really represent only one variable species. The 
name of it must therefore be Mimomys savini Hinton, which is the oldest valid name. 
In fact all possible intermediate forms between typical M. intermedius, M. savini 
and M. majori are present in the abundant material from the Upper Freshwater 
Bed (Pasquier 1972). The presence of three species of one genus with identical 
dimensions in one and the same layer is also scarcely imaginable from the ecological 
point of view. Specimens determined as 'M. majori and 'M. savini' have also been 
found in some Czechoslovak and German fossil localities, always associated with 
M. intermedius. 

Genus ARVICOLA Lacepede 1799 

The systematic study of even the living representatives of this genus presents 
many difficulties to zoologists. Miller (1912) divided living populations of water 
voles from western Europe into seven species but later investigators (for bibliography 
see Reichstein 1963) have demonstrated that nearly all the characters which were 
used for the diagnosis of these species lie within the limits of individual variation. 
Variations occur principally in the dimensions and in some proportions of the skull, 
proportions which may change allometrically with the changes of absolute dimensions. 

Cytological studies support the view that there are only two living species in 
this genus, A. sapidus Miller 1908, from the south of France and the Iberian Penin- 
sula, and A. terrestris (Linnaeus 1758) distributed throughout the rest of Europe 


as well as the northern part of Asia and the Middle East. The Recent population 
in Britain belongs to this last species. Reichstein (1963), after a very thorough 
analysis, found distinct but small craniological differences between the two species 
of Arvicola and further differences were reported by Corbet et al. (1970). 

The systematic study of fossil forms, usually possible only from teeth and frag- 
mentary skulls, is still more difficult. Remains from glacial deposits are usually 
larger than those from the interglacials and interstadials or from the Holocene, and 
most of the characters which have been used for creating new fossil species of water 
voles are expressions of individual variability, present also in Recent species. It 
seems beyond doubt that Arvicola developed from a late form of the genus Mimomys, 
most likely Mimomys savini, through the progressive reduction of the formation of 
roots in the molars and acquisition of the continual growth of these teeth. One 
character, typical of Mimomys, is nevertheless retained in geologically older popu- 
lations of Arvicola : the enamel of the cheek-teeth is thinner on the concave and 
thicker on the convex sides of the salient angles, the reverse being true in the late 
Pleistocene and Recent populations. The two forms are united by a full series of 
intermediate specimens. O. Fejfar (in Koenigswald 1970) re-examined the holotype 
of Mimomys cantianus Hinton from the Hoxnian deposits of Ingress Vale, Swans- 
combe, and found that there is no trace of root-formation. This form must therefore 
be included in the genus Arvicola. Arvicola cantiana (Hinton) is the oldest available 
name for the primitive representatives of Arvicola and later names such as A. 
hactonensis, A . greeni and A . praeceptor must be treated as synonyms. 

Koenigswald (1972, 1973) also studied the stratigraphical range of the various 
stages of the Mimomys- Arvicola lineage from European localities and defined a 
series of faunal groups : 

(a) A Mimomys savini fauna, including the Upper Freshwater Bed of West Runton. 

(b) and (c) Arvicola faunas groups i and 2, both with A. cantiana (Koenigswald 
included Swanscombe in group 2). 

(d) Arvicola fauna group 3, with A. terrestris. 





Fig. 19. Development of the enamel layer in Middle European forms ot Arvicola. Arvicola 
cantiana (Elster-Riss) ; Arvicola cantiana-terrestris transition form (Riss-Eem) ; 
Arvicola terrestris (Eem-Holocene) . From Koenigswald (1973 : 164). 

























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Mimomys savir 

/ fauna 

Arvicola fauna 

group 1 

Arvicola fauna, g 

roup 2 


A, fauna, 

gp 3 

Fig. 20. Chronological distribution of Arvicola and other stratigraphically important 
small mammal species in European Pleistocene faunas. From Koenigswald (1973 : 161). 

He referred the Glutton Stratum of Tornewton Cave to an intermediate stage 
between groups 2 and 3 (Fig. 20). 

Koenigswald made a further study of British Arvicola remains in 1974 and (pers. 
comm.) assigned the phylogenetic position, between A. cantiana and A. terresiris, 
of the finds from various sites as follows : 

A. cantiana 


> intermed 


late forms 


— >A. terrestns 



Joint Mitnor Cave 

Kent's Cavern 

(Barnfield Pit and 

(Cave Earth) 

Ingress Vale) 


Swanton Morley 

Wye Cave 



Tornewton Cave 
(Otter Stratum) 

( = Merlin's Cave) 


Tornewton Cave 

Tornewton Cave 

(Reindeer Stratum) 

(Glutton Stratum) 

These findings are of very far-reaching stratigraphical importance. A. cantiana 
appears for the first time in Britain at the 'late Cromerian' site of Westbury-sub- 
Mendip, which appears to represent Koenigswald's group i, and is also present 
in the Hoxnian deposits of Clacton and Swanscombe. The transition form is 
represented by remains from the cold stage represented by the Glutton Stratum of 


Tornewton Cave, in the Otter Stratum of that cave (where it is apparently slightly 
more advanced) and in the Last Interglacial hippopotamus faunas of Barrington, 
Joint Mitnor Cave and Swanton Morley. A. terrestris is characteristic of Last 
Glaciation and Holocene faunas. 

It must be noted that Hinton's and Koenigswald's divisions between A . cantiana 
and A. terrestris do not exactly coincide. Koenigswald, from a re-examination of 
the specimens from the Barrington hippopotamus site, considered that the Arvicola 
remains from this locality should be attributed to the A . cantiana-terrestris transition 
form rather than to A. praeceptor {= A. cantiana), as determined by Hinton (1926b : 
393). Stuart (see p. 69) has suggested that both forms are present. The specimens 
from the Last Interglacial site of Hessle, Yorkshire, attributed by Hinton (in Bisat 
1940) to A. praeceptor, but which Boylan (1967) considered to lie within the normal 
range of variability of A. terrestris, have not been re-examined during the present 

Surprisingly the remains from Aveley, Grays Thurrock, Harkstead and Stutton 
(currently widely regarded as Last Interglacial, though most if not all are sites without 
hippopotamus - the undoubted hippopotamus remains from Grays were collected 
more than a century ago and their exact provenance within the terrace complex of 
that area is unknown) are advanced A. cantiana and appear to be earlier than the 
pre-hippopotamus Glutton Stratum of Tornewton Cave. 

The Cray ford record of A . cantiana must be accepted with some reserve. Although 
Arvicola is recorded from this locality by Dawkins & Sanford (1866) and by Whitaker 
(1889) it was not mentioned by Hinton (1900a, b, 1910b, 1926b). The present 
determination is based on a single specimen in the British Museum (Natural History). 

Arvicola cantiana (Hinton 1910) 
Water vole, extinct 

1846 Arvicola amphibia ; Owen : 201-205 (partim). 

1882a Arvicola amphibius ? Linnaeus ; Newton : 87-88. 

1902 Microtus intermedins ; Hinton & White : 414-415. 

1910b Mimomys cantianus Hinton : 491. 

1926b Arvicola bactonensis Hinton : 386-389. 

1926b Arvicola greeni Hinton : 389-391, fig. 106, 1-2. 

1926b Arvicola praeceptor Hinton : 391-394, fig. 106, 3-17. 

1970 Arvicola cantiana (Hinton) ; Koenigswald : 418-420. 

Localities : Ostend, Norfolk (57) : Owen 1846, Newton 1882a, Lydekker 1885, 
Hinton 1926b, BM(NH). 

Westbury-sub-Mendip, Somerset (13) : Bishop 1974, 1975. 

Clacton-on-Sea, Essex (43) : Hinton in Warren 1955, Sutcliffe 1964, BM(NH). 

*Swanscombe (Ingress Vale and Barnfield Pit, Lower Loam), Kent (30) : Hinton 
& White 1902, Stopes 1904, Hinton 1910b, 1926b, Barrett-Hamilton & Hinton 
1910-21, Sutcliffe 1964, Koenigswald 1970, BM(NH). 

Aveley, Essex (34) : BM(NH). 



Grays Thurrock, Essex (33) : Hinton, Kennard & Newton 1900, Hinton 1910b, 
1926b, Barrett-Hamilton & Hinton 1910-21, BM(NH). 

Ilford, Essex (35) : Dawkins & Sanford 1866, Whitaker 1889, Newton 1890a, 
Johnson & White 1900, Hinton 1900a, b, 1910b, 1926b, BM(NH). 

Stutton, Suffolk (44) : Stuart collection and seen in Ipswich Museum. 

Harkstead, Suffolk (44) : Ipswich Museum - Stuart {in litt.). 

Northfleet, Kent (29) : det. A. Stuart. 

Crayford, Kent (27) : Dawkins & Sanford 1866, Whitaker 1889, BM(NH). 

Arvicola cantiana-terrestris transition form 
(see pp. 99-102 above) 
Localities : Tornewton Cave (Glutton Stratum and Otter Stratum), Devon (3) 
Kowalski 1967, (BM(NH). 

Barrington, Cambridgeshire (40) : Hinton 1926b (but see note on p. 69), BM(NH). 
Joint Mitnor Cave (layer IV), Devon (i) : BM(NH). 
Minchin Hole, Glamorganshire (16) : det. A. J. Stuart. 
Stoke Tunnel Beds, Suffolk (46) : Ipswich Museum. 
Swanton Morley, Norfolk (66) : Stuart, pers. comm. 

Fig. 21. Distribution of fossil remains of 
Arvicola cantiana (Hinton) (■) and A. 
terrestris (Linn.) (D) in the British Isles. 
(Transition form WJ, not studied #.) 

Arvicola terrestris (Linnaeus 1758) 
Water vole 

1846 Arvicola amphibia ; Owen: 201-20^ (partim). 

1847 Hypudaeus spelaeus Cuv. ; Giebel : 88. 

1872 Arvicola amphibius Desm. ; Dawkins & Sanford : 180. 

1910a Arvicola abbotti Hinton : 34-35. 

i960 Arvicola amphibius terrestris Bramwell : [10]. 


Localities : Ightham Fissures, Kent (28) : Newton 1894, Hinton 1910a, b, 
Barrett-Hamilton & Hinton 1910-21, Hinton 1926b, Zimmermann 1959, BM(NH). 

Kent's Cavern (Cave Earth), Devon (6) : Owen 1846, Dawkins & Sanford 1866, 
Blackmore & Alston 1874, Lydekker 1885, Hinton 1926b, Kennard 1945-46, 
Carreck 1957, BM(NH). 

Tornewton Cave (Reindeer Stratum), Devon (3) : Kowalski 1967. 

Wye (= Merlin's) Cave, Herefordshire (18) : Bate 1901, Hinton 1926b, BM(NH). 

Arvicola cantiana / A. terrestris complex 
(phylogenetic position not studied) 

With the exception of Hoxne (Hoxnian) and Bacon Hole, Hessle, Kirkdale Cave 
and the Hyaena Stratum of Tornewton Cave (which are Last Interglacial) the 
following sites are all believed to be of Last Glaciation or Holocene age. The 
Arvicola remains are probably mostly advanced forms. 

Localities : Hoxne, Suffolk (54) : Wolff {in Hit.). 

Bacon Hole (16) : Stuart {in Hit.). 

Bobbitshole, Suffolk (46) : Ipswich Museum, det. J. N. Carreck. 

Hessle, Yorkshire (74) : Lamplugh 1891, Bisat 1940, Catt & Penny 1966, Boylan 
1967, specimen in Institute of Geological Sciences Museum, London, not seen 
by K. K. 

Kirkdale Cave, Yorkshire {yy) : Owen 1846, Giebel 1847, Dawkins & Sanford 
1866, Lydekker 1885, Hinton 1926b, BM(NH). 

Tornewton Cave (Hyaena Stratum), Devon (3) : Kowalski 1967. 

Aveline's Hole, Somerset (11) : Davies 1921, Hinton 1921, 1924, 1926b, BM(NH). 

Brixham Cave, Devon (4) : Hinton 1926b, Carreck 1957, BM(NH). 

Chudleigh Fissure, Devon (7) : BM(NH). 

Dog Holes Cave, Lancashire (79) : Jackson 1910, 1912, Hinton 1926b, BM(NH). 

Elder Bush Cave, Staffordshire (69) : Bramwell 1964. 

Gough's Cave, Somerset (12) : BM(NH). 

Langwith Cave, Derbyshire (72) : Mullins 1913, BM(NH). 

Levaton Cave (2) : Carreck 1957. 

Pin Hole Cave, Derbyshire (73) : Jackson 1934, 1947. 

Happaway Cave, Devon (5) : Hinton 1926b, Carreck 1957, BM(NH). 

Nazeing, Essex (37) : Hinton 1952, BM(NH). 

Cowside Cave No. 3, Yorkshire (78) : BM(NH). 

Thatcham, Berkshire (Holocene, 23) : Wymer 1962. 

General distribution of Arvicola, and systematic remarks. As stated above, 
the forms described as A. greeni, A. bactonensis and A. pr acceptor must be treated 
as synonyms of A. cantiana. The first two of the above-mentioned species were 
described by Hinton (1926b) from specimens collected at Ostend, near Bacton, 
Norfolk, during the first part of the nineteenth century. A. bactonensis is known 


only from the holotype and A. greeni is represented only by two isolated teeth. 
The stratigraphic position of these remains is uncertain. Hinton (1926b) suggested 
that they were contemporary with the Cromer Forest Bed or only slightly younger. 
As Arvicola is not represented at the Cromerian type locality of West Runton, where 
the ancestral form Mimomys savini is present, it must be supposed that these 
specimens are from later deposits. They may be equivalent in age to the faunal 
assemblage of Westbury-sub-Mendip, where A. cantiana is present. 

A. cantiana is known, under various names, from numerous localities of Holsteinian 
(Hoxnian) age in continental Europe and Siberia. Remains of A. terrestris, to 
which species it gave rise, are abundant from the Upper Pleistocene of Europe, 
including England and Wales. Like other voles, Arvicola never reached Ireland. 

A. terrestris is now widely distributed in nearly all Europe and in cold and tem- 
perate parts of Asia, as far south as Persia and Palestine. It lives mostly near water, 
but under diverse climatic and vegetational conditions, ranging from the taiga belt, 
through deciduous forest and steppe, to the deserts in the south. As a species of 
high ecological tolerance it was present during the Pleistocene under both glacial 
and interglacial conditions. 

Genus PITYMYS McMurtrie 1831 

Many zoologists are of the opinion that Pitymys is only a subgenus of Microtus. 
Pitymys is morphologically similar to Microtus, sensu stricto, and is rather similar in 
its ecology, but the living forms are biologically quite different. Chaline (1972), 
who studied fossil vole remains from the French Pleistocene, came to the conclusion 
that fossil forms of Pitymys are nothing but morphotypes of Allophaiomys plio- 
caenicus Kormos 1932. If this is correct, then A. pliocaenicus would be a junior 
synonym of Pitymys arvaloides and its name must be changed accordingly. In his 
opinion the Recent subgenus Pitymys is polyphyletic and its members are more 
closely related to different forms of Microtus than to one another. In the opinion 
of one of us (K. K.) Chaline underestimates the ecological similarity of Recent 
species of Pitymys and all other zoological evidence, basing his conclusions on the 
pattern of M^ only. For this reason he is unable to follow Chaline's suggestions. 

Voles of the Pitymys group dominated the European vole faunule during the 
Cromerian Interglacial. They were still common during the Hoxnian, after which 
they disappeared completely from Britain. Their distribution in Europe and 
western Asia is now rather discontinuous. The first evidence of the presence of 
Pitymys in Britain was published by Major in 1902. Hinton (1923a) described two 
different species from the Upper Freshwater Bed of West Runton, P. gregaloides and 
P. arvaloides. They are clearly distinct species, though in continental Europe 
intermediate specimens are present. If Pitymys is only a subgenus of Microtus, a 
very prevalent opinion among Recent mammalogists, the name Pitymys arvaloides 
Hinton is a taxonomic homonym and must be replaced by a new name, Microtus 
{Pitymys) arvalidens Kretzoi 1958. In the opinion of K. K. it is more convenient 


to treat Pitymys as a separate genus and the name given by Hinton is therefore 
retained here. 

Pitymys arvaloides Hinton 1923 
Pine vole, extinct 

1882a Arvicola avvalis Pallas ; Newton : 88-89, pl- xiv, figs 2-5. 
1902 Microtus {Pitymys) sp. ; Major : 107, fig. 15 (28). 

1923a Pitymys arvaloides Hinton : 541-542. 

1958 Microtius {Pitymys) avvalidens Kretzoi : 57 (new name for Pitymys arvaloides Hinton 
1923, non Arvicola arvaloides Pomel). 

1972 Allophaiomys pliocaenicus Kormos ; Chaline : 104. 

Localities : West Runton (Upper Freshwater Bed), Norfolk (65) : Newton 
1882a, Major 1902, Hinton 1923a, 1926b, Kretzoi 1958, Chaline 1972, BM(NH). 
Hitchin, Hertfordshire (39) : Carreck 1959 : 326 (det. as P. cf. arvaloides). 
Swanscombe (Barnfield Pit, Lower Gravel and Lower Loam), Kent (30) : Carreck 

1959 (: 326), Sutcliffe 1964. 

Fig. 22. Distribution of remains of Pitymys 
arvaloides Hinton in the British Isles. 

Distribution in the British Isles. P. arvaloides is abundant in the Cromerian 
Upper Freshwater Bed at West Runton. It is present also in deposits of the 
lower part of the Hoxnian Interglacial, but disappears completely before its end. 
There is at present no evidence of its presence in Britain during the Anglian cold 
phase and it is probable that it invaded this country from the European continent 

General distribution and Systematic remarks. P. arvaloides is present at 
numerous localities of 'Cromerian' age on the continent (Italy, Yugoslavia, Germany, 
Hungary, Poland, Czechoslovakia, Austria, the European part of the Soviet Union 
and Georgia). Nearly everywhere it has been found together with P. gregaloides. 
It is not impossible that the two species, different in the morphology of M^ but of 
identical dimensions, are only two forms of one polytypic species. 


Pity my s gregaloides Hinton 1923 
Pine vole, extinct 

1882a Arvicola gregalis Pallas ; Newton : 90-91, pi. xiv, figs 6, 6a. 

1 89 1 Microtus {Arvicola) gregalis Pallas ; Newton : 53. 

1923a Pitymys gregaloides Hinton : 541-542. 

1972 Allophaiomys pliocaenicus Kormos ; Chaline : 104. 

Localities : West Runton (Upper Freshwater Bed), Norfolk (65) : Newton 
1882a, 1891, Hinton 1923a, 1926b, BM(NH). 
Westbury-sub-Mendip Fissure, Somerset (13) : Bishop 1974, 1975. 
Kent's Cavern, Devon (6) : Hinton 1926b, Kennard 1945-46, BM(NH). 

Fig. 23. Distribution of remains of Pitymys 
gregaloides Hinton in the British Isles. 

Distribution in the British Isles, P. gregaloides is present in Cromerian 
deposits at West Runton and at the Westbury Fissure, together with P. arvaloides. 
Two specimens, one preserved in the British Museum (Natural History), the other 
in Bristol Museum, are known from Kent's Cavern. The stratigraphical position of 
these specimens is unknown, but the species has not yet been recorded from any 
British Hoxnian locality and it is likely that they are of pre-Hoxnian age. 

General distribution and Systematic remarks. P. gregaloides is present, 
together with P. arvaloides, at numerous localities in central and southern Europe 
dated as Cromerian {sensu lato) . 

Genus MICROTUS Schrank 1798 

The genus Microtus now holds the dominant position in the subfamily Microtinae, 
being both most highly differentiated and most individually numerous. In the 
fossil state molar teeth make up most of the finds, mandibles occurring less commonly 
and complete skulls only in exceptional cases. Furthermore, it should not be 
forgotten that in living populations the variability of the molar pattern is sometimes 



enormous (Kowalski 1970) and this variability may be discontinuous, representing 
genetical polymorphism. At the same time there are species, clearly differentiated 
in external characters and in biological requirements, in which the teeth are nearly 
identical (e.g. the lower teeth of M. agrestis and M. arvalis cannot be clearly dis- 
tinguished, nor can the upper teeth of M. arvalis and M. oeconomus). All these 
factors make the correct determination of fossil remains, including the material 
from Britain described below, very difficult. 

The genus Microtus appeared rather late in the European Pleistocene, being 
found first in the Cromerian Interglacial when Pitymys was still numerically domi- 
nant. In the Hoxnian Interglacial both genera lived together but in later sediments 
in Britain Pitymys disappeared completely and on the European continent it now 
plays only a subordinate role. 

Microtus arvalinus Hinton 1923 
Vole, extinct 

1902 Microtus sp., recalling M. arvalis ; Major : 107, fig. 15 (27). 

1923a Microtus arvalinus Hinton : 541-542. 

1972 Microtus [Microtus) arvalis (Pallas) ; Chaline : 106. 

Localities : West Runton (Upper Freshwater Bed), Norfolk (65) : Major 1902, 
Hinton 1923a, 1926b, BM(NH). 

Westbury-sub-Mendip, Somerset (13) : Bishop (pers. comm.). 

Swanscombe (Lower Loam and Upper Middle Gravel), Kent (30) : Schreuder 
1950, Sutcliffe 1964, BM(NH). 

Fig. 24. Distribution of remains of Microtus 
arvalinus Hinton in the British Isles. 

General distribution. M. arvalinus was present in Britain during the 
Cromerian and Hoxnian interglacials and the intervening Westbury stage. A small 
form of Microtus, with the M^ pattern of the M. arvalis] agrestis group, is present in 


the Cromerian deposits of France, Germany, Austria, Czechoslovakia, Hungary, 
Romania and Yugoslavia. 

Systematic remarks. M. arvalinus is identical in its M^ pattern with Recent 
representatives of the M. arvalisjagrestis group, and differs from these only in its 
slightly smaller dimensions. It is probable that M. arvalinus is the ancestral form 
of M. agrestis. According to Chaline (1972), M. arvalinus is a synonym of M. 
arvalis, but there does not seem to be any good reason for putting it into this 
synonymy rather than that of M. agrestis. 

Microtus agrestis (Linnaeus 1761) 
Field vole 

1846 Ayvicola agrestis ; Owen : 206-207. 

1847 Hypudaeus hucklandii Giebel : 88. 

1894 Microtus (= Arvicola) agrestis ; Newton : 197. 

1910b Microtus agrestis neglectus ; Hinton : 494. 
1910b Microtus agrestoides Hinton : 493. 

Microtus arvalis and Microtus arvalisjagrestis group (partim) ; many authors. 

General notes on the Microtus arvalis I agrestis group. During the nineteenth 
century most zoologists did not appreciate the differences between M. arvalis and 
M. agrestis and these two names must be regarded as synonyms in papers published 
at that time. Later on, the presence of the M. arvalis group on the Orkney Islands 
{M. arvalis does not survive today on the mainland of Britain and M. agrestis does 
not occur on Orkney) led to the hypothesis that M. arvalis was the first Microtus 
to reach Britain in the postglacial period. Under the influence of this hypothesis 
fossil remains of the M. arvalisjagrestis group were usually determined as 'M. 
arvalis'. It must nevertheless be born in mind that only the upper M^ provides 
diagnostic characters for distinguishing M. agrestis and M. arvalis. Both lower 
jaws and other isolated teeth are useless for this purpose (Dienske 1969). An 
example of the confusion which can occur in this context is provided by the Microtus 
remains from Dog Holes Cave, Lancashire. All the upper dentitions were correctly 
determined as 'M. agrestis', but all the lower jaws were labelled 'M. corneri' (= M. 
arvalis) . This illustrates well the tendency to decide in each doubtful case in favour 
of M. arvalis. 

In the site list for M. agrestis given below all the localities of M. agrestis, 'M. 
agrestoides' , 'M. arvalis I agrestis group' and 'M. arvalis' are enumerated. 

Localities : Clacton-on-Sea, Essex (43) : Hinton 1923b, Singer et at. 1973. 

Swanscombe (Upper Middle Gravels), Kent (30) : Schreuder 1950 {M. arvalis/ 
agrestis group). 

Ilford (Uphall Estate), Essex (35) : Hinton 1900a, b, Johnson & White 1900. 

Grays Thurrock, Essex (33) : Hinton, Kennard & Newton 1900, Hinton 1901, 
1910b, Barrett-Hamilton & Hinton 1910-21, Hinton 1924, 1926b, BM(NH) {'M. 


Harkstead, Suffolk (44) : Ipswich Museum. 

Stutton, Suffolk (44) : Ipswich Museum. 

Northfleet, Kent (29) : Burchell 1935. 

Tornewton Cave (Glutton Stratum, Hyaena Stratum, Reindeer Stratum, Dilu- 
vium), Devon (3) : Kowalski 1967, BM(NH). 

Water Hall Farm Pit, Hertfordshire (38) : BM(NH). 

Alveston Fissure, Gloucestershire (15) : det. G. B. Corbet. 

Bacon Hole, Glamorganshire (16) : det. A. J. Stuart. 

Bobbitshole, Suffolk (46) : Ipswich Museum, det. J. N. Carreck. 

Barrington, Cambridgeshire (40) : BM(NH). 

Joint Mitnor Cave, Devon (i) : BM(NH). 

Kirkdale Cave, Yorkshire (77) : Owen 1846, Giebel 1847, Dawkins & Sanford 
1866, Blackmore & Alston 1874, BM(NH). 

Minchin Hole, Glamorganshire (16) : det. A. J. Stuart. 

Aveline's Hole, Somerset (11) : Davies 1921, Hinton 1921, 1924, BM(NH). 

Bleadon Cave, Somerset (10) : BM(NH). 

Brixham Cave, Devon (4) : BM(NH). 

Chudleigh Fissure, Devon (7) : BM(NH). 

Dog Holes Cave, Lancashire (79) : Jackson 1910, 1912, BM(NH). 

Elder Bush Cave, Staffordshire (69) : Bramwell 1964. 

Gough's Cave, Somerset (12) : BM(NH). 

Great Doward Cave, Herefordshire (17) : BM(NH). 

Ightham Fissures, Kent (28) : Newton 1894, 1899b, Hinton 1910b, Barrett-Hamil- 
ton & Hinton 1910-21, Hinton 1921, 1926b, Jackson 1947, Zimmermann 1959. 

Kent's Cavern, Devon (6) : Owen 1846, Dawkins & Sanford 1866, Blackmore & 
Alston 1874, Lydekker 1885-87, Kennard 1944-45, BM(NH). 

King Arthur's Cave, Herefordshire (17) : BM(NH). 

Langwith Cave, Derbyshire (72) : Mullins 1913, BM(NH). 

Levaton Cave, Devon (2) : Carreck 1957. 

Merlin's Cave, Herefordshire (18) : Hinton 1925. 

Pin Hole Cave, Derbyshire {y^) : Jackson 1934, 1947. 

Brean Down, Somerset (9) : Apsimon, Donovan & Taylor 1961. 

Happaway Cave, Devon (5) : BM(NH). 

Lynx Cave, Denbighshire (67) : Blore 1966. 

Cowside Cave No. 3, Yorkshire (78) : BM(NH). 

Corstorphine, nr. Edinburgh, Scotland (81) : Evans 1913. 

Creag nan Uamh Cave, Inchnadamph, Scotland {8^) : Peach & Home 1917. 

Distribution in the British Isles. M. agrestis is now widely distributed in 
Great Britain and many of the remains cited as fossils are probably of relatively 
recent age, in any case post-glacial. 

The supposed earliest British occurrence of this group or species of rodents is at 
the Hoxnian locality of Swanscombe, where it is recorded as having been found in 
the Upper Middle Gravel in association with M. arvalinus (Schreuder 1950). The 
occurrence together of these two forms, which, it has been suggested (p. 109), 
are members of the same phylogenetic lineage, creates a problem. Schreuder 


Fig. 25. Distribution of fossil remains of 
the Microtus arvalis/agrestis group in the 
British Isles. 

distinguished the two forms at Swanscombe on grounds of size and stated that she 
would not have hesitated to accept as M. arvalinus the specimen determined by her 
as 'Microtus sp., arvalisfagrestis group' had it not been so large. It has been pointed 
out by Corbet, however (pers. comm.), that there is great variation, which may 
reflect the age of the animal concerned, in the crown size of teeth of any microtine 
in which the teeth continue to grow during life. In view of the very small size of 
the sample studied by Schreuder, the occurrence of both M. arvalinus and M. arvalis/ 
agrestis together in the same deposit is accepted here with some reserve. The 
remains are stratigraphically placed at about the time of transition between these 
two forms and a larger collection of remains is needed for study. 

M. agrestis was common during the warm stages represented by both Grays 
Thurrock and Joint Mitnor Cave, and in the Glutton Stratum of Tornewton Cave, 
but is absent from the faunas of Cray ford and Clevedon Cave. It was common 
during the Last Glaciation. At most localities where fossil remains have been found 
only lower jaws were present, but in each case where upper teeth were also found the 
second upper molar is of the typical M. agrestis pattern. There can be no doubt 
that the field vole was discontinuously present in Britain from at least the time of the 
Grays Thurrock deposits. 

Hinton (1910b) created the name 'M. agrestoides for the specimens of M. agrestis 
from Grays Thurrock. He was surely influenced here by the supposition that Grays 
Thurrock has a very ancient fauna. The only distinctive character of his new 
species is 'the constant development of a fourth outer angle in the last upper molar'. 
This 'fourth angle' is present in a great proportion of specimens of Recent M. 
agrestis and its presence is not sufficient for specific determination. 

According to Jackson (1929b), a skull of a vole which he regarded as close to M. 
arvalis was found in Kilgreany Cave, Co. Waterford, Ireland. This is the only 
possible evidence that voles ever reached Ireland ; it was a skull with all teeth 
missing. It was found in the 'Lower Stalagmite' together with Dicrostonyx and 
human remains ; '. . , a close examination of the sockets suggests that the species 


is the continental field vole Microtus arvalis or a close ally, and not the common field 
vole {M. agrestis)' (Jackson 1929b : 147). 

The specimen has recently been re-examined by Savage (1966), who noted that 
it is notably fresh and unaltered and who had no doubt that it should be referred to 
M. agrestis. He pointed out that occasional vole remains have been found in north- 
east Ireland in pellets dropped by passing owls. On the basis of existing evidence, 
it is unlikely that voles ever reached Ireland. 

General distribution. M. agrestis is now widely distributed in Europe, as 
well as in the cold and temperate parts of Asia. In Europe it extends north to 
include the whole Scandinavian peninsula and Finland. Fossil remains are known 
in continental Europe beginning with the Riss (penultimate) Glaciation. 

Microtus arvalis (Pallas 1779) 
Common vole 

1910a Microtus corneri Hinton : 35-36. 

Microtus arvalis [partim) ; many authors. 

Distribution in the British Isles. Hinton (in Barrett-Hamilton & Hinton 
1910-21, 2 : 467) correctly stated that : 

'On several occasions fossils from various British deposits of Late Pliocene and Pleistocene age 
have been determined as belonging to M. arvalis (e.g. from fissures near Bath, Somerset, by 
Blackmore and Alston, Proc. Zool. Soc, 1874, 468) ; but in most cases such records imply, 
because of the fragmentary nature of the material, on which they are based, nothing more 
than the presence of a "vole" with an arvaloid, i.e. a normal dentition.' 

According to Hinton [loc. cit.) the cranial remains from the Ightham fissures are 
'apparently identical' with M. arvalis ; this locality probably marks the date of the 
arrival of the species in Britain. 

Hinton (1910a) described from the Ightham fissures a new species, M. corneri, 
very similar to the Orkney vole, M. orcadensis. We now know, however, that M. 
orcadensis is nothing other than an isolated population of M. arvalis, possibly 
developed within historic times. 

M. corneri has been recorded from the following localities : 

Aveline's Hole, Somerset (11) : Davies 1921, Hinton 1921, 1924. 

Brixham Cave, Devon (4) : BM(NH) (det. Hinton). 

Dog Holes, Lancashire (79) : BM(NH) (det. Hinton). 

Ightham Fissures, Kent (28) : Hinton 1910a, b, Barrett-Hamilton & Hinton 
1910-21, Hinton 1921, 1926b, Jackson 1929, Hinton 1952, Zimmermann 1959. 

Langwith Cave, Derbyshire (72) : Hinton 1910a, MuUins 1913. 

Levaton Cave, Devon (2) : Carreck 1957 {^Microtus arvalis group, cf. corneri 

The status of M. arvalis, including M. corneri, m the British Pleistocene requires 
extensive further study. Although known today on the Orkney Islands, M. 
arvalis does not occur on the British mainland. There is geological evidence that 


the Orkney Islands have never been connected to the mainland since the last glacial 
phase, when ice covered the British Isles as far south as Norfolk and South Wales, 
and the local population of M. arvalis [= orcadensis] must therefore be a recent 
introduction from the continent (Corbet 1961). In Europe, M. arvalis is typical of 
the zone of mild climate, being absent from the Scandinavian peninsula and from 
northern Denmark. Its distribution as well as ecological data make its presence 
in the British Isles in the early postglacial period extremely improbable. The 
majority of fossil records of 'M. corneri' from Britain are of rather doubtful value. 
As mentioned above, lower jaws of Microtus (which are specifically undeterminable) 
from Dog Holes Cave have been identified as M. corneri whilst all the skulls belong, 
without exception, to M. agrestis. The skull from Brixham Cave determined as 
belonging to M. corneri is probably a specimen of M. oeconomus. 

One of the most potentially important collections for the study of M. arvalis is 
that from Ightham Fissures, Kent, where many excellently preserved vole skulls 
were collected. It is evident, however, that remains of both Pleistocene (Last 
Glaciation) and Holocene age have been mixed together. In such circumstances it 
is possible that sometimes a single species is represented in the fossil material by 
two different subspecies or by two different populations of different dimensions. 
Among the Microtus material from this locality, besides the typical skulls of M. 
gregalis and M. agrestis, there are two groups of larger and smaller skulls. Some of 
them may belong to M. oeconomus, and the skull selected as the holotype of M. 
corneri probably belongs here. Other specimens could be young individuals of this 
species, or they may really belong to M. arvalis. 

Recently two small further collections of rodents have been studied by Corbet 
(pers. comm.) which suggest that M. arvalis may indeed have been present in Eng- 
land during the Last Glaciation. These are from Marlow, Buckinghamshire (25, 
Treacher Collection), and Beckford, Worcestershire (19, Briggs and Coope Collection), 
all in the British Museum (Natural History) . The Marlow remains were apparently 
predominantly M. arvalis. There was also one first lower molar of M. oeconomus. 
Corbet considers that the lack of any second upper molars of M. agrestis, and the 
occurrence of only one lower molar of M. oeconomus among the many first lower 
molars (the teeth which distinguish these last two closely related species from M. 
arvalis), suggest that M. arvalis is the principal species present. The age of the 
Marlow rodents is uncertain, but is likely to be Last Glaciation. 

Corbet considers that the Beckford specimens, if all one species, must be M. 
arvalis. They could be a mixture of M. agrestis and M. oeconomus, but the absence 
of the diagnostic second upper molar of M. agrestis or the first lower molar of M. 
oeconomus makes this improbable. Beckford is of Last Glaciation age, with a cold 
insect-fauna and ^*C dates of 27 650 ± 250 (Birmingham 293) and 27 300 ± 500 years 
(Birmingham 595). 

At the present state of our knowledge neither the status of M. arvalis in the 
British Pleistocene nor the systematic position of 'M. corneri' is clear. Craniometric 
analysis of the whole collection of Microtus from Ightham would be of great value. 
If M. arvalis was present in the British Isles it was an invasion of short duration 
during one of the phases of the Last Glaciation. 


Microtus nivaloides Major 1902 
Vole, extinct 

1902 Microtus nivaloides Major : 106, fig. 19. 

1923a Microtus nivalinus Hinton : 541-542. 

Locality : West Runton (Upper Freshwater Bed), Norfolk (65) : Major 1902, 
Barrett-Hamilton & Hinton 1910-21, Hinton 1923a, 1926b. 

Fig. 26. Distribution of remains of Microtus 
nivaloides Major in the British Isles. 

Distribution in the British Isles. M. nivaloides is present in the Cromerian 
of Norfolk. 

General distribution. Small representatives of the genus Microtus with the 
Mj resembling that of M. nivalis, determined as M. nivalinus Hinton, M. nivaloides 
Major or M. suhnivalis Pasa, have been recorded in 'Cromerian' deposits in Germany, 
Czechoslovakia, Hungary, Romania, Yugoslavia, Italy and the Ukraine. 

Systematic remarks. In addition to the well-defined form M. nivaloides, 
another from the same layer and locality was described as M. nivalinus by Hinton 
(1923a). Intermediate specimens between these two forms also occur and it is 
probable that only one species is represented. M. nivaloides, widely distributed in 
deposits of 'Cromerian' age in Europe, is probably the ancestor of M. nivalis, a 
common species in late Pleistocene localities. In Chaline's opinion (1972 : 151) 
M. nivaloides and M. nivalinus are synonyms of M. nivalis. This is in contradiction 
to the picture of evolution in the M. nivalis line as presented by the same author 
who considered that M. nivalis developed from M. malei not earlier than the 
penultimate glaciation. 

Microtus nivalis (Martins 1842) 
Snow vole 

1907a Microtus nivalis (Martins) ; 
1907a Microtus malei Hinton : 49 

Hinton : 39-48 ; pi. i, figs 5-23- 
pi. I, figs 24-27. 


Localities : Crayford and Erith (27) : Hinton 1907a, b, 1910b, Barrett-Hamilton 
& Hinton 1910-21, Hinton 1926b, Kennard 1944, Jackson 1947, Chaline 1972, 

Clevedon Cave, Somerset (14) : Hinton 1907a, b, 1910b, Barrett-Hamilton & 
Hinton 1910-21, Hinton 1926b, Chaline 1972, BM(NH). 

Cow Cave, Chudleigh (7) : det. G. B. Corbet. 

Gough's Cave, Somerset (12) : BM(NH). 

Tornewton Cave (Glutton Stratum), Devon (3) : Kowalski 1967, BM(NH). 

Water Hall Farm Gravel Pit, Hertfordshire (38) : BM(NH). 

East Wickham, Plumpstead, London : Hinton 1907b, 1926b, BM(NH). 

Some additional records of M. nivalis exist in early papers. This species has 
been mentioned from a cave in the Forest of Dean, from Fisherton, Grays Thurrock 
and Swanscombe. Hinton (1926b) did not mention these localities in his monograph 
as containing M. nivalis and no remains of the snow vole have been seen by the 
writers among material collected from these sites. 

Fig. 27. Distribution of fossil remains of 
Microtus nivalis (Martins) in the British 

Distribution in the British Isles. M. nivalis seems to be a typical element 
of the fauna of the period represented by Crayford and the Tornewton Cave Glutton 
Stratum. There was a stage when the genus Microtus was represented in the 
British Isles by two species only, M. nivalis and M. oeconomus. It is entirely 
absent from Devensian localities in Britain. 

General distribution. M. nivalis is now distributed in mountain ranges, 
mostly in the Mediterranean area, from the Pyrenees in the west to Lebanon, the 
Caucasus and Kopet-Dag in the east. It inhabits treeless, mostly rocky or stony 
localities, not necessarily cold ones. During the Pleistocene it was more widely 

Systematic remarks. M. nivalis has a very variable tooth-pattern. Its 
presence in Britain is beyond doubt, but the existence of a second species from this 
group, M. malei, does not seem probable. The teeth determined by Hinton (1907a) 
as belonging to M. malei were found exclusively in localities where M. nivalis is 


numerous, and all intermediate forms between typical nivalis and typical malei can 
be found. M. malei must therefore be taken as a synonym of M. nivalis. But 
according to Chaline (1972) M. malei is a valid species, and he designates as its 
lectotype the specimen M26481 in the British Museum (Natural History) from 
Clevedon Cave. In his opinion all the specimens from Clevedon Cave, determined 
by Hinton as belonging to M. nivalis, M. malei and M. ratticeps, represent only one 
variable species, M. malei. M. nivalis developed, according to Chaline, from M. 
malei during the late Pleistocene and was not present in Britain. His hypothesis, 
as stated above, seems to be contradictory, but it is possible that the representatives 
of the group of M. nivalis present in the lowland of Europe, including Britain, were 
different from the Recent M. nivalis from the European mountains. In such a case 
they must be named Microtus malei Hinton. 

Microtus ratticepoides Hinton 1923 
Vole, extinct 
1923a Microtus ratticepoides Hinton : 541-542. 

Localities : West Runton (Upper Freshwater Bed), Norfolk (65) : Hinton 
1923a, 1926b, BM(NH). 

Swanscombe (Lower Loam and Upper Middle Gravel), Kent (30) : Schreuder 1950, 
Sutcliffe 1964, BM(NH). 

Fig. 28. Distribution of remains of Microtus 
ratticepoides Hinton in the British Isles. 

Distribution in the British Isles. M. ratticepoides is known from deposits of 
Cromerian and Hoxnian age. 

General distribution. M. ratticepoides has been recorded, mostly in 
'Cromerian' faunal assemblages, in Holland, Germany, Romania, Czechoslovakia, 
Poland, Hungary and the European part of the Soviet Union including Georgia. 

Systematic remarks. The tooth-pattern of M. ratticepoides is nearly identical 
to that of M. oeconomus, but the dimensions of the fossil form are slightly smaller 


than those of the Recent one. M. ratticepoides may be the ancestor of the Middle 
Pleistocene and Recent M. oeconomus. 

Microtus oeconomus (Pallas 1776) 
Root vole 

1870b Arvicola ratticeps (Blasius) ; Sanford : 124-125, pi. viii, figs i, la-id. 
1890a Microtus {Arvicola) ratticeps Key. and Bl. ; Newton : 453-456, figs 1-2, 
1 90 1 Microtus ratticeps ; Bate : 103-104. 

Localities : Crayford and Erith (27) : Whitaker 1889, Newton 1890a, Hinton 
1910b, Barrett-Hamilton & Hinton 1910-21, Hinton 1926b, Kennard 1944, Jackson 
1947, Hinton 1952, BM(NH). 

Harkstead, Suffolk (44) : Ipswich Museum. 

Clevedon Cave, Somerset (14) : Hinton 1910b, Barrett-Hamilton & Hinton 
1910-21, BM(NH). 

Tornewton Cave (Glutton Stratum, Reindeer Stratum, Diluvium), Devon (3) : 
Kowalski 1967, BM(NH). 

Bacon Hole, Glamorganshire (16) : det. A. J. Stuart. 

Water Hall Farm Pit, Hertfordshire (38) : BM(NH). 

Aveline's Hole, Somerset (11) : Davies 1921, Hinton 1921, 1924, BM(NH). 

Bleadon Cave, Somerset (10) : Palmer 1934, BM(NH). 

Chudleigh Fissure, Devon (7) : BM(NH). 

Dog Holes Cave, Lancashire (79) : Jackson 1910, Barrett-Hamilton & Hinton 
1910-21, BM(NH). 

Elder Bush Cave, Staffordshire (69) : Bramwell 1964. 

Fisherton, Wiltshire (22) : Blackmore & Alston 1874, Bate 1901, Jackson 1910, 
Barrett-Hamilton & Hinton 1910-21, BM(NH). 

Gough's Cave, Somerset (12) : BM(NH). 

Great Doward Cave, Herefordshire (17) : BM(NH). 

Ightham Fissures, Kent (28) : Newton 1894, Bate 1901, Jackson 1910, Hinton 
1910b, Barrett-Hamilton & Hinton 1910-21, Hinton 1926b, Zimmermann 1959, 

Isleworth, Middlesex (26) : BM(NH). 

Kent's Cavern, Devon (6) : Sanford 1870b, Kennard 1945-46, BM(NH). 

King Arthur's Cave, Herefordshire (17) : BM(NH). 

Langwith Cave, Derbyshire (72) : Barrett-Hamilton & Hinton 1910-21, Mullins 


Levaton Cave, Devon (2) : Carreck 1957. 

Marlow, Buckinghamshire (25) : BM(NH). 

Merlin's Cave, Herefordshire (18) : Hinton 1924. 

Picken's Hole, Somerset (10) : Stuart 1974. 

Pin Hole Cave, Derbyshire (73) : Jackson 1934, 1947. 

Dowel Cave, Derbyshire (71) : Bramwell i960. 

Etches' Cave, Derbyshire (71) : Pernetta 1966. 


Lynx Cave, Denbighshire (67) : Blore 1966. 

Nazeing, Essex (37) : Hinton 1952, BM(NH). 

Cowside Cave No. 3, Yorkshire (78) : BM(NH). 

Hay Wood Rockshelter, Somerset (10) : BM(NH). 

Somerset levels, Huntspill Cut (8) : Hinton 1952. 

Nornour, Isles of Scilly (91) : Pernetta & Handford 1970. 

Creag nan Uamh Cave, Sutherland (83) : Peach & Home 1917. 

Fig. 29. Distribution of fossil remains of 
Microtus oeconomus (Pallas) in the British 

Distribution in the British Isles. M. oeconomus is a widely distributed fossil 
species in England and Wales, which also reached northern Scotland. It is a 
dominant element of the Crayford fauna and is numerous in the sediments of the 
Last Glaciation. It was still present at the end of this period (Nazeing) and dis- 
appeared during postglacial times. 

General distribution. M. oeconomus is now widely distributed in the northern 
parts of Europe, Asia and North America. In Europe its westernmost localities 
are in Holland and Germany and isolated colonies are known in Hungary and 
Austria. Fossil remains are common in the Late Pleistocene and Holocene layers 
of central Europe including France, Hungary and Switzerland. 

Systematic remarks. In Chaline's opinion (1972) M. ratticepoides is a synonym 
of M. oeconomus, which developed during the Last Glaciation, its ancestor being 
M. malei. This hypothesis is rather contradictory and lacks evidence, at least as 
far as material from the British Isles is concerned. 

Microtus gregalis (Pallas 1779) 
Narrow-skulled vole 

1894 Microtus (= Arvicola) gregalis ; Newton : 197-198, pi. xi, fig. 12. 

1910a Microtus anglicus Hinton : 36-37. 

Localities : Aveline's Hole, Somerset (11) : Davies 1921, Hinton 1921, 1924, 


Bleadon Cave, Somerset (10) : BM(NH). 

Brixham Cave, Devon (4) : BM(NH). 

Chudleigh Fissure, Devon (7) : BM(NH). 

Great Doward Cave, Herefordshire (17) : BM(NH). 

Ightham Fissures, Kent (28) : Newton 1894, Hinton 1910a, b, Barrett-Hamilton 
& Hinton 1910-21, Hinton 1924, 1926b, 1952, Zimmermann 1959, Chaline 1972, 

Isleworth, Middlesex (26) : BM(NH). 

Kent's Cavern^ Devon (6) : Kennard 1945-46, BM(NH). 

King Arthur's Cave, Herefordshire (17) : BM(NH). 

Langwith Cave, Derbyshire (72) : Hinton 1924, Mullins 1913. 

Levaton Cave, Devon (2) : Carreck 1957 {'Microtus cf. anglicus'). 

Merlin's Cave, Herefordshire (18) : Hinton 1925. 

Picken's Hole, Somerset (10) : Stuart 1974. 

Pin Hole Cave, Derbyshire (73) : Jackson 1934, 1947. 

Tornewton Cave (Reindeer Stratum), Devon (3) : Kowalski 1967, BM(NH). 

Uphill Cave, Somerset (9) : Hinton 1926b. 

Dowel Cave, Derbyshire (71) : Bramwell i960. 

Happaway Cave, Devon (5) : BM(NH). 

Nazeing, Essex (37) : Hinton 1952, BM(NH). 

Fig. 30. Distribution of fossil remains of 
Microtus gregalis (Pallas) in the British 

Although M. anglicus ( = M. gregalis) is recorded from Dog Holes Cave (Jackson 
1912) four specimens from this locality preserved in the British Museum (Natural 
History) and labelled by Hinton as M. anglicus do not belong to this species but to 
M. agrestis. 

Distribution in the British Isles. M. gregalis was present in the southern 
parts of England during the Last Glaciation, but is absent in all earlier sediments. 
It probably reached Britain only once and disappeared before the end of the last 
glacial period. 


General distribution. M. gregalis, the unique representative of the subgenus 
Stenocranius Kastschenko, is now distributed widely in northern and central Asia 
and in arctic Europe from the White Sea in the west to the Bering Strait in the east. 
In the western parts of its range it is associated with the tundra belt but in the east, 
where the climate is more continental, it is present also in the steppe and desert 
zone in eastern and central Asia. 

In Europe it was common in the time of the Last Glaciation, reaching France and 
Britain in the west and Switzerland and Hungary in the south. According to 
Chaline (1972), it was present in France during both the last and penultimate 

Systematic remarks. M. gregalis was recorded in the Pleistocene of Europe 
by Nehring (1875 : 7). According to Hinton (1924), fossil remains from Europe 
are specifically distinct from M. gregalis and approach M. tianschanicus Biichner 
(now regarded as synonym of M. eversmanni Poljakov). In the modern systematics 
of this group M. eversmanni is recognized as a subspecies of M. gregalis. The 
morphology and dimensions of the European fossil specimens lie completely within 
the range of variability of the polytypic species M. gregalis. According to Chaline 
(1972) the name anglicus can be preserved to designate the subspecies of M. gregalis 
from the Last Glaciation localities of Europe. 

Genus LAGURUS Gloger 1841 

Lagurus lagurus (Pallas 1773) 

Steppe lemming 

1967 Lagurus lagurus (Pallas) ; Kowalski : 115 -119, figs 1-24. 

Fig. 31. Distribution of fossil remains of 
Lagurus lagurus (Pallas) in the British 

Distribution in the British Isles. Remains of Lagurus lagurus have been 
found at only one locality, in the Glutton Stratum of Tornewton Cave, Devon (3) 
(Kowalski 1967). Sixty-three first lower molars (3-8 per cent of the total number of 


rodent remains in this layer) and many other teeth were found in this deposit, which 
dates from the cold stage preceding the interglacial hippopotamus-bearing Hyaena 
Stratum of this cave. 

General distribution. L. lagurus developed during the Middle Pleistocene 
from more primitive forms of the genus Lagurus. It is now distributed in the steppe 
zone of Eurasia from the River Dnepr to Mongolia and Chinese Turkestan. Its 
fossil remains are relatively common in Upper Pleistocene localities in central and 
eastern Europe, but less so in western Europe. In France (Chaline 1973) it has 
been found in strata of the penultimate ('Riss') and early last (Wiirm) glaciations. 

Systematic remarks. The fossil British Lagurus remains are identical in 
dimensions and molar pattern with recent specimens of this species. The only 
character not observed in contemporary material is a tendency among the Tornewton 
specimens towards the fusion of the two anterior enamel triangles on the first lower 
molar (Kowalski 1967). 

IV. history of the rodent fauna of the BRITISH ISLES DURING 


In the earlier parts of this work rodent localities and rodent species were con- 
sidered individually. In conclusion, let us attempt to construct from this infor- 
mation a generalized table of the rodent faunas of the various stages of the British 
Pleistocene. The construction of such a table must nevertheless be carried out with 
reserve. Rodent faunas were not uniform throughout the British Isles. Cold- 
adapted species in the north were able to coexist with more temperate species in the 
south, and during every glacial and interglacial stage there occurred a shifting of the 
vegetational zones which would have caused corresponding movements of the rodent 

Sometimes sea barriers prevented the movement of rodents. The known rodent 
faunas of the Lower Pleistocene and of the later glaciations of England are similar 
to those of the continent of Europe, suggesting land connections, whereas only a 
few rodent species are known from the later interglacials, suggesting separation. 
There is no evidence that any rodents crossed to Ireland until some stage during the 
Last Glaciation. 

Fossil rodent material is seldom rich enough to demonstrate local or temporal 
differentiation within particular divisions of the Pleistocene of the British Isles and 
there are some stages, especially the earlier glaciations, from which no rodent remains 
are yet known. The incompleteness of the record of British rodents was strikingly 
demonstrated by the recent discovery of previously unrecognized rodent faunas in 
the Westbury Fissure and in Tornewton Cave. Further discoveries can be expected 
to provide an ever more detailed sequence for a long time to come. 

The most likely stratigraphical relationship of the principal British rodent localities, 
at our present incomplete state of knowledge, and the known range of the various 
rodent species, is shown in Table 12. A series of faunal stages can be recognized. 
In ascending order these are as follows. 


Red Crag. The study of rodents from the eariiest Pleistocene deposit of the 
British Isles, the Red Crag, is very difficult because of the mixing in this marine 
deposit of contemporary and derived fossils. Castor fiber is apparently a con- 
temporary species. Trogontherium minus, known from Pliocene sediments on the 
continent of Europe, and Hystrix are apparently derived remains of pre-Pleistocene 

ICENiAN Crag and Pastonian Forest Bed. The next faunal assemblage is that 
of the Icenian Crag and the earlier (Pastonian) part of the Forest Bed Series at 
East Runton. Mimomys pliocaenicus (from both deposits) and M. reidi (from the 
Icenian Crag) are species of great stratigraphic importance. On the continent of 
Europe they are limited to the Upper Villafranchian or Tiglian. There is no evidence 
of any difference between the British and European continental rodent faunas at this 
stage. Trogontherium hoisvilletti and Castor fiber are two long-ranging rodents also 
present in both series of deposits. 

Cromerian sensu stricto. The rodent fauna from the Cromerian type locality at 
West Runton is unusually rich, there being at least eleven species. Trogontherium 
boisvilletti and Castor fiber reappear. Mimomys savini replaces M. pliocaenicus ; 
Apodemus sylvaticus, Cricetus cricetus, Clethrionomys glareolus, Pitymys arvaloides, 
P. gregaloides, Microtus arvalinus, M. nivaloides and M. ratticepoides make an 
appearance. Sciurus whitei occurs in the overlying marine 'Monkey Gravel'. 

The West Runton fauna is predominantly of forest and meadow type, testifying 
to a mild climate, though Cricetus cricetus could point to colder or more continental 
conditions. No arctic elements are present. Allophaiomys pliocaenicus and lem- 
mings, already present on the continent at the time of the earliest glaciations, are 
so far unrecorded from West Runton. 

'Westbury stage'. The rich mammalian fauna from Westbury Fissure, described 
by Bishop (1974), does not at present fit into the generally accepted Pleistocene 
sequence of the British Isles. Bishop has presented evidence of a previously un- 
recognized stage later than the Cromerian sensu stricto of West Runton and possibly 
equivalent to Mosbach and other continental sites. 

Although the richest rodent-bearing level at Westbury (the 'Rodent Earth') is 
the latest deposit in the fissure and may be slightly younger than the main bone 
deposit (the 'Calcareous Series') five rodent species are nevertheless represented in 
the latter. Apodemus sylvaticus, Pitymys gregaloides and Microtus arvalinus 
reappear ; Arvicola cantiana and Lemmus sp. appear for the first time. All these 
forms persist into the 'Rodent Earth' which also contained Dicrostonyx sp., the 
earliest record in the British Pleistocene, and the only record from the British 
Pleistocene of Pliomys episcopalis. Clethrionomys glareolus reappears. Of special 
interest is the first appearance in the British Pleistocene of Arvicola cantiana, 
which apparently replaces Mimomys savini, believed to be an earlier stage in the 
same phylogenetic lineage, and which is so common at West Runton. 

Castor fiber is represented in the lowest water-laid deposits of the cave. The 
lowest deposits of Kent's Cavern and the 'Forest Bed' deposit of Ostend, Norfolk, 
may also belong to this stage. 

chronological distribution of fossil r 




Tabie I 

t localities and rodent species fi 



:. 9,500 years ago), Thalcham 

e, Cow Cave (part), 7Goughs Cave (part), 7Hutton Cave 




Ha,™d ,e.<( 

Ha, Wood) 






















! 1 




I 1 1 

1 i 1 

1 1 1 




? 1 
















JoinI Milnor 
Cave inlarglacial 


1 1 





— 1 — u 



1 1 








""group 1"™ 



Arv/co/a launa 


1 ' 1 












! 1 


. , >' 









1 1 






MuNDESLEY ARCTIC Bed. The Spermophilus described by Newton (1882b) from 
a deposit with remains of arctic plants at Mundesley, Norfolk, provides the only 
known instance of a fossil rodent from a British Lower or Middle Pleistocene non- 
interglacial deposit. According to Newton the deposit was immediately overlain 
by thick glacial deposits which West & Wilson (1966) attributed to the Lowest of tian 
(Anglian) Glaciation. They referred underlying interglacial deposits to the 
Cromerian sensu stricto. 

For the time being the exact relationship between the Mundesley Arctic Bed and 
the Westbury deposits must remain uncertain. If, as might be expected, Mundesley 
is later than Westbury then it is necessary to explain why the Westbury stage has 
not been recognized, except possibly at Ostend, in the Forest Bed sequence of East 
Anglia. If, on the other hand, it is earlier then it is necessary to postulate a hitherto 
unrecognized cold stage between the Cromerian sensu stricto and the 'Westbury' stage. 

HoxNiAN DEPOSITS. The best sequence of deposits with rodent remains of 
Hoxnian age is that of Barnfield Pit, Swanscombe. A considerable time interval is 
believed to have occurred within this sequence, between the Lower Loam and the 
Middle Gravel (see p. 52). From the lowest deposit, the Lower Gravel, only 
Pitymys arvaloides has been recorded. In the overlying Lower Loam this species 
is accompanied by Castor fiber, Arvicola cantiana, Microtus arvalinus and M. 

These last two species continue into the Upper Middle Gravel, where they are 
joined by Clethrionomys glareolus and Lemmus sp. Although Trogontherium 
hoisvilletti and Apodemus sylvaticus have not been recorded from Barnfield Pit, they 
are known from a nearby pit in the same terrace deposits at Ingress Vale. Trogon- 
therium hoisvilletti appears to have been common during the Hoxnian, being recorded 
also from Clacton, Copford and Hoxne. It is unknown from later deposits. 

Comparison of the rodent faunas of West Runton, Westbury-sub-Mendip and 
Swanscombe. Although the geological evidence indicates at least one major glaci- 
ation between the time of accumulation of the deposits at West Runton and at 
Swanscombe, and although there are very great differences between the large 
mammals of the three sites mentioned above, changes in the rodent fauna were 
remarkably gradual. The distribution of the various rodent species is shown in 
Table 13. 

Whilst incomplete collecting probably accounts for many of the gaps in the table 
a gradual replacement of species is nevertheless apparent. Mimomys savini, 
abundant at West Runton, appears to be replaced at Westbury by Arvicola cantiana. 
This species persists at Swanscombe and in slightly later deposits. Microtus 
arvalinus and M. ratticepoides are present at both West Runton and Swanscombe 
(the former occurs also at Westbury), but apparently disappear from the British 
Pleistocene after the Upper Middle Gravel. Trogontherium hoisvilletti and Pitymys 
arvaloides are of similar distribution, last seen at Ingress Vale and in the Lower 
Loam of Barnfield Pit respectively. Pitymys gregaloides occurs at West Runton 
and Westbury but does not reappear at Swanscombe. Microtus nivaloides is not at 
present known from deposits later than West Runton. Pliomys is at present known 



Table 13 

Comparison of rodent species represented at West Runton, 
Westbury-sub-Mendip (Bishop 1974 ^^^ pers. comm.) and Swanscombe 











Barnfield Pit 


















Sciurus whitei 


Trogontherimn boisvilletti 



Castor fiber 




Apodemus sylvaticus 





Cricetus cricetus 


Dicrostonyx sp. 


Lemmus sp. 




Clethrionomys glareolus 




Pliomys episcopalis 


Mimomys savini 


Arvicola cantiana 





Pitymys arvaloides 




P. gregaloides 




Microtus arvalinus 






M. nivaloides 


M. ratticepoides 




only from the youngest deposit at Westbury. Dicrostonyx and Lemmus, so common 
in British Upper Pleistocene deposits, appear for the first time at Westbury. Lemmus 
reappears in the Upper Middle Gravel of Swanscombe. 

No major break is discernible in the above sequence of rodent faunas. The 
Westbury fauna has some characters in common with both West Runton and 
Swanscombe, confirming Bishop's supposition that it is of intermediate age. The 
rodent faunas of the three localities are similar to Koenigswald's Mimomys savini 
fauna, Arvicola fauna group i and Arvicola fauna group 2 of the continent of Europe 
(see p. loi) and suggest land connections during the periods concerned. A similarity 
of the large mammals of Swanscombe and those from the Holsteinian site of Steinheim 
an der Murr in Germany has previously been observed by Sutcliffe (1964). 

Notes on the Wolstonian-Ipswichian part of the succession. Although 
palaeobotanical evidence provides support for only one further interglacial in the 


British Isles after the Hoxnian it is difficult to fit all the known Upper Pleistocene 
rodent faunas into such a simple sequence of events. The generally accepted 
Wolstonian-Ipswichian part of the succession appears, from the rodent and other 
evidence previously described, to be composite. The so-called Tpswichian' inter- 
glacial is here interpreted as double, with the Middle Terrace of the Thames rep- 
resenting a warm stage earlier than the Joint Mitnor Cave warm stage. 

Middle Terrace of the Thames interglacial. Reasons for regarding the 
deposits of the Middle Terrace of the Thames (type locality Grays Thurrock) as 
representing the earlier of these two warm periods have been given on pp. 55-58. 
Ilford, Aveley, Harkstead, and Stutton in Suffolk are also included here. The 
known rodent fauna of this stage is very sparse. Arvicola cantiana reappears ; 
other species include Castor fiber, Apodemus sylvaticus, Clethrionomys glareolus and 
Microtus agrestis. Consideration needs to be given to the possibility that this 
stage is Hoxnian (Zeuner (1945) believed that Grays was of Penultimate Interglacial 
= Hoxnian age) but the occurrence of Arvicola cantiana in other Middle Terrace 
localities such as Aveley and Stutton, both of which have been claimed on palaeo- 
botanical grounds to be Ipswichian, makes it unnecessary to resort to such an early 

ToRNEWTON Cave Glutton Stratum cold stage. The Arvicola from the 
Glutton Stratum of Tornewton Cave is the transition form between A . cantiana and 
A. terrestris, suggesting that this deposit is later than the Middle Terrace of the 
Thames where the species is A. cantiana. We know that this stage was followed by 
a further interglacial, since these deposits were immediately overlain by the Hyaena 
Stratum, with remains of hippopotamus. Other species from the Glutton Stratum 
are Microtus oeconomus and M. nivalis, which make up the bulk of the rodent fauna, 
and also M. agrestis, Lagurus lagurus (the only British record of this species), cf. 
Allocricetus bursae and Cricetus cricetus. Dicrostonyx torquatus and Lemmus lemmus 
are present, but not abundant. 

This assemblage closely resembles that of the Late Middle Terrace of Crayford 
and Erith, regarded by Hinton (1926b) as later than that of the Middle Terrace of 
Grays Thurrock. Rodents common to both Crayford and the Tornewton Cave 
Glutton Stratum are Microtus oeconomus, M. nivalis, M. agrestis, Arvicola sp., 
Lemmus lemmus and Dicrostonyx torquatus. Most of the species that are not com- 
mon to both deposits are so rare in the British Pleistocene that stratigraphic com- 
parison with other localities cannot be made. Spermophilus primigenius from 
Crayford and Cricetus cricetus from Tornewton Cave are unknown from any other 
post-Hoxnian British sites ; cf. Allocricetus bursae (Tornewton Cave) is known from 
only one other British locality, Hutton Cave, which is of uncertain age. Lagurus 
lagurus is known only from Tornewton Cave. 

The mollusc evidence suggests that the Corbicula Bed at Crayford, in which most 
of the rodent remains were found, is still interglacial, though absence of forest 
species of land mollusca indicates open grassland. The living representatives of the 
subgenus Urocitellus, to which Spermophilus primigenius belongs, are also open- 
country species. On the continent 5. primigenius is unknown later than the 
'penultimate' glaciation. 


From the available evidence the most likely stratigraphic position for Crayford 
would seem to be the end of the Grays-Ilford-Aveley interglacial. 5. primigenius 
prevents us from attributing a date late in the Pleistocene to the Crayford deposits. 
Crayford apparently heralds the arrival of the Tornewton Cave Glutton Stratum 
rodent fauna and marks the beginning of the arrival of a great wave of eastern 
species of rodents from the European continent. Characteristic of this particular 
invasion of continental forms are Spermophilus primigenius, Cricetus cricetus, 
Allocricetiis bursae, Lagurus lagurus and Microtus nivalis. In addition, there 
arrived three other boreal species, Dicrostonyx torquatus, Lemmus lemmus and 
Microtus oeconormis. During the coldest phase of this period all rodents connected 
with forest environment, including such adaptable species as Clethrionomys glareolus 
and Apodemus sylvaticus, apparently disappeared from the British Isles. The 
absence of Microtus gregalis, so common in Last Glaciation deposits, is a characteristic 
feature of the fauna of this stage. 

The apparent occurrence of this cold stage between two interglacial stages with 
Tpswichian' floras is not in accordance with current palaeobotanical opinion, which 
identifies only one Ipswichian. The severity of this cold stage requires further 
assessment. The occurrence of both reindeer and wolverine, in addition to the 
above-mentioned rodent species, suggests more than a minor cool phase within an 
interglacial. Intense disturbance of the Glutton Stratum and the mixing of it 
with a vast quantity of fragments of broken stalagmite formations could be inter- 
preted as evidence of very intense frost disturbance later than the Glutton Stratum 
but earlier than the deposition of the overlying layers. The relationship of this 
apparent cold stage to known glacial stages of the Upper Pleistocene (ice of the 
penultimate glaciation is believed to have reached north Devon) must remain, for 
the time being, unknown. 

Other localities which Hinton (1926b) considered, from the rodent evidence, to be 
contemporaneous with Crayford are Clevedon and Banwell Caves. Cow Cave (part), 
Gough's Cave (part), Northfieet and Water Hall Farm (each with Microtus nivalis), 
and Hutton Cave (with cf. Allocricetus bursae) may be further examples of deposits 
approximately contemporaneous with the Glutton Stratum of Tornewton Cave. 

Microtus nivalis apparently disappears from the British Pleistocene after this 

Joint Mitnor interglacial. The occurrence of a further interglacial stage with 
hippopotamus is reliably demonstrated at Tornewton Cave where the interglacial 
Hyaena Stratum overlies the Glutton Stratum. The occurrence of remains of M. 
nivalis and M. oeconomus, apparently underlying hippopotamus-bearing deposits, 
at Water Hall Farm gravel pit provides a further example of this relationship. 

Important hippopotamus sites dating from this interglacial probably include 
Barrington, Joint Mitnor Cave and Swanton Morley. The rodent fauna of this 
stage is very sparse. The Arvicola cantiana-terrestris transition form and Microtus 
agrestis are the most common species. Clethrionomys glareolus, Apodemus sylvaticus 
and Castor fiber also occur. The Otter Stratum of Tornewton Cave (with the clawless 
otter, Cyrnaonyx, and white-toothed shrew, Crocidura, unknown elsewhere in the 
British Pleistocene) may represent the beginning of this stage, before the arrival of 


hippopotamus ; Minchin Hole appears to represent the end, after it had disappeared. 
The sparseness of this fauna suggests isolation of the British Isles from the European 
continent at that time. 

Before leaving discussion of this interglacia-l, represented also by other sites such 
as Trafalgar Square with plant remains, we must refer once more to the Middle 
Terrace of the Thames interglacial, indistinguishable from it on palaeobotonical 
evidence. Evidence from the rodent remains nevertheless makes it difficult to 
explain the sequence other than with two post-Hoxnian interglacial stages separated 
by the Tornewton Glutton Stratum cold stage. The sequence becomes even more 
complicated if we consider the significance of the Baker's Hole cold stage. Was this 
the same as the Tornewton Cave Glutton Stratum cold stage or (if, as Carreck {in 
litt.) has suggested, it occurred between the Ilford and Crayford stages) does it 
represent yet another earlier cold stage within the Tpswichian', sensu lato? 

There is at present no means of knowing to which of these warm stages the 
Ipswichian type locality of Bobbitshole belongs, although recent studies by Coope 
(1974) have showed very close similarities between the insects of this site and of 
Trafalgar Square, suggesting that it could be the later one. This is in agreement 
with the view of Evans (1971) who assigned the Ipswichian sensu stricto to his half- 
cycle 3W (see pp. 40-42). 

Last Glaciation. The Last (Devensian) Glaciation opened the way for a new 
immigration. This time the immigrants included not only both species of lemmings 
but also two new arrivals, the ground squirrel Spermophilus superciliosus and the 
vole Microtus gregalis. Cricetus, Lagurus and AUocricetus, though present at this 
time in France, did not find their way back to Britain. Arvicola had by this time 
evolved to A. terrestris (Koenigswald's Arvicola fauna group 3, see pp. loo-ioi). 
Apodemus sylvaticus, Clethrionomys glareohts and Microtus agrestis are other species 
recorded and M. arvalis may have been present for a short time. Castor fiber 
apparently disappeared. 

This fauna differs substantially from that of the preceding cold phase represented 
by the Glutton Stratum of Tornewton Cave, from which it is readily distinguishable. 
Diagnostic criteria include the absence of Spermophilus primigenius (found at 
Crayford), Cricetus, AUocricetus, Microtus nivalis and Lagurus and the appearance of 
Spermophilus superciliosus and Microtus gregalis. Arvicola had evolved from the 
A. cantiana-terrestris transition form to true A. terrestris. Dicrostonyx, Lemmus and 
Microtus oeconomus were present during both stages. 

It is impossible at present to reconstruct in detail all the changes of the rodent 
fauna of the Last Glaciation. We have nevertheless now reached the most recent 
stage within the Quaternary, except for the Holocene, and the later part of it lies 
within the range of radiocarbon dating. Some changes of the rodent fauna are 
apparent, which offer a basis for more detailed study in the future, when more and 
better-dated material becomes available for study. 

During the coldest part of the Last Glaciation Apodemus sylvaticus and perhaps 
also Clethrionomys glareolus disappeared from a large part or perhaps all of Britain. 
During the warmest interstadial the two lemmings were apparently absent, at least 
in the south of England. 


Three sites with i*C dates are of special importance. These are : 

1. Willment's Pit, Isleworth, west London. 43 140+1520 or —1280 years b.p. 
Microtus oeconomus and M. gregalis (possibly the earliest record of this species in 
Britain) occur ; no lemmings. 

2. Upton Warren, Worcestershire. 41 000 years b.p. Dicrostonyx present. 

3. Beckford, Worcestershire. 27 650 + 250 years b.p. Probably with Microtus 

Coope, who is making a detailed study of the insect faunas of the Last Glaciation, 
has named the period of time covered by the above sites the Upton Warren Inter- 
stadial complex. He found evidence (pers. comm.) of an intense and short amelior- 
ation of climate at about 43 000 years ago, followed by climatic deterioration with 
increase of continentality at about 41 000 years. The lack of lemmings at Isleworth 
is in accordance with the evidence provided by the insects. The Beckford insect 
fauna (pers. comm.) indicates cold conditions. 

Other important Last Glaciation sites in the south of England, which lack 
lemmings, are Levaton Cave and Picken's Hole. The deposits of this last-mentioned 
site are interpreted by Stuart (1974) as possibly Early Devensian (layer 5, with 
Microtus cf. oeconomus and M. cf. gregalis) and Middle Devensian (layer 3, with 
Citellus sp, (= Spermophihis sp.) and Microtus cf. gregalis). 

The best evidence of the rodent faunas of the end of the Last Glaciation and 
early Holocene is provided by the deposits of the Lea Valley and by the Peakland 
Archaeological Society's excavations in Dowel Cave, Etches' Cave and Fox Hole 
Cave. The end of the Pleistocene is marked by a special abundance of lemmings. 
Late Glacial deposits in the Lea Valley (dated on palaeobotanical evidence) contained 
both Lemmus lemmus and Dicrostonyx torquatus and also Arvicola terrestris, M. 
oeconomus and M. anglicus (= M. gregalis). Of these Dicrostonyx torquatus was 
still present in pollen Zone HI (the last stage of the Pleistocene, ending about 
10 000 years ago) and M. oeconomus survived into the Holocene. The cave sites 
mentioned above also indicate an abundance of lemmings in the Late Pleistocene. 
Other species present were Arvicola terrestris, M. gregalis and M. oeconomus, the 
last-mentioned species surviving there too into the Holocene. 

At the end of the Last Glaciation arctic species of rodents began to disappear, 
though both Lemmus and Dicrostonyx seem to have survived at Nazeing until 
near the end of this period, the latter species still being present in pollen Zone IH. 
M. gregalis does not appear to have persisted into post-Pleistocene times. 

Holocene. The postglacial rodent fauna is poorer than that of the neighbouring 
continent of Europe, but two species of rodents, now extinct in the British Isles, 
persisted. These are Microtus oeconomus and Castor fiber. M. oeconomus apparently 
continued in the Lea Valley until pollen Zone V, Mesolithic, about 9000 years ago. 
It also survived until Mesolithic times at Dowel Cave and was apparently present on 
the Isles of Scilly during the Bronze Age. 

Castor fiber, apparently absent during the Last Glaciation, had already reappeared 
by about 9500 years ago (pollen Zone IV-V transition, Mesolithic, Star Carr), 
surviving until possibly the thirteenth century. Although the date of the Scottish 
finds is uncertain, Castor fiber may have reached Scotland during this stage. 


Arvicola terrestris persisted from the Pleistocene and still occurs in Britain at the 
present day. In the Lea Valley Apodemus sylvaticus, Clethrionomys glareolus and 
Microtus agrestis reappeared in pollen Zone V-VI (about gooo years ago) after an 
apparent interval at the end of the Pleistocene. 

Scmrus vulgaris was probably a post-Pleistocene arrival. Others were Apodemus 
Jiavicollis, Muscardinus avellanarius and Micromys minutus. It is difficult to decide 
whether these last three species reached Britain with or without participation of 
human activity. Geological and ecological data suggest rather the second pos- 
sibility. Glis glis, Mus musculus, Rdttus rattus and Rattus norvegicus are evidently 
recent human introductions. 

Ireland. At some stage during the Last Glaciation the two species of lemmings, 
but no voles nor any steppe elements, managed to cross to Ireland. A ^*C date of 
33 000 years for an associated mammoth bone from Castlepook Cave suggests that 
Lemmus and Dicrostonyx were probably established in Ireland at that time. Both 
are northern species which might be expected to reach Ireland in advance of other 
rodents. On the basis of existing evidence, the availability of lemmings for immigra- 
tion from the British mainland appears entirely plausible. 

Apodemus sylvaticus, recorded from the Pleistocene levels of Castlepook Cave, 
may also have arrived in Ireland during the Pleistocene, although it has been 
observed that at some cave sites this species was most abundant in the upper levels, 
suggesting a later date. Other rodent species which occur in Ireland today are 
probably post-Pleistocene arrivals. 


The writers gratefully acknowledge help with the compilation of this Bulletin 
from many sources. 

Dr W. von Koenigswald, of Tubingen University, advised on British representa- 
tives of the Mimomys- Arvicola lineage, and commented on the typescript. 

Mr M. J. Bishop, of University College, London, provided us with full details of 
his determinations of the rodent remains from the Middle Pleistocene fissure deposit 
at Westbury-sub-Mendip and allowed us to see the typescript of his report on this 
site before publication. 

Mr J. N. Carreck (Queen Mary College) read the typescript and provided many 
valuable suggestions. He also allowed us to mention important findings from his 
own unpublished studies in the Thames estuary. 

Dr G. B. Corbet, of the British Museum (Natural History), provided determinations 
of rodent remains from Alveston Fissure, Beckford and Marlow and commented on 
the typescript. 

Dr A. J. Stuart, of the University of Cambridge, provided determinations of 
rodent remains excavated in Minchin Hole in 1973 and much other help. He 
commented on our typescript and allowed us to see that of his review of British 
Pleistocene vertebrates before publication. 


Dr J. d'A. Waechter provided an important collection of rodent remains excavated 
from the Lower Loam of Barnfield Pit, Swanscombe, during 1968-72. 

Dr G. R. Coope, of Birmingham University, provided rodent remains from Beckford 
and advice on the events of the Last Glaciation in Britain. 

Dr R. G. Wolff, of the University of Florida, provided details of rodent remains 
collected from Hoxne, Norfolk, by the University of Chicago under grants GS41435 
and GS2907 of the National Science Foundation to Dr R. Singer. 

Stratigraphic and other information was received from Mr D. Bramwell (Peakland 
Archaeological Society), Dr A. S. Clarke (Royal Scottish Museum, Edinburgh), 
Dr J. Jewell (British Museum (Natural History)), Mr D. Mayhew (Cambridge), 
Professor G. F. Mitchell (Trinity College, Dublin), Mr H. E. P. Spencer (Ipswich), 
Dr R. G. West and Mrs G. Wilson (University of Cambridge). 

Mr J. J. Hooker sieved Pleistocene sediment from Aveley, Essex, for rodent 
remains, Mr J. Simons sediment from Willments Gravel Pit, Isleworth, London, and 
from Water Hall Farm Pit, Hertfordshire, all with important results. 

Mr A. P. Currant (British Museum (Natural History)) prepared Figs i, 5-18 and 
21-31 and Mrs A. J. Sutcliffe drew Figs 2-4. 

Mr D. L. F. Sealy provided much editorial help, including preparing the index, 
and Mrs M. N, M. Bhukhureea prepared the typescript with great patience. 


An asterisk (*) distinguishes those papers cited in the text which are not directly concerned 
with Quaternary rodents of the British Isles. 

Abbot, W. J. L. 1917. The ossiferous fissures of the valley of the Shode. Appendix I in : 
Bennett, F. J. Ightham, The Story of a Kentish Village and its Surroundings : 101-119, 
I pi., 2 tables, I fig. London. 

* Agadzhanian, a. K. 1973. (Collared lemmings of the Pleistocene). Nov. tekt. nov. 

otlozhi i chelovek, Moscow, 5 : 320-355. [In Russian.] 
Apsimon, a. M., Donovan, D. T. & Taylor, H. 1961. The stratigraphy and archaeology 
of the late-glacial and post-glacial deposits at Brean Down, Somerset. Proc. speleol. Soc, 
Bristol, 9 : 67-136. 

* AzzAROLi, A. 1953. The deer of the Weybourne Crag and Forest Bed of Norfolk. Bull. Br. 

Mus. nat. Hist. (Geol.), London, 2 : 1-96. 
Barrett-Hamilton, G. E. H. & Hinton, M. A. C. 1910-21. A History of British Mammals 

(unfinished). 1 (1910) : xvi-l-263 pp., 20 pis. 2 (191 1 -21) : 748 pp., 37 pis. London. 
Bate, D. M. A. 1901. A short account of a bone cave in the Carboniferous Limestone of the 

Wye Valley. Geol. Mag., London, (n.s. 4) 8 : 101-106. 
Beirne, B. p. 1947. The history of the British Land Mammals. Ann. Mag. nat. Hist., 

London, (11) 14:501-514. 
BiSAT, W. S. 1940. Older and Newer Drift in East Yorkshire. Proc. Yorks. geol. Soc, 

Leeds, 24 : 137-151. 
Bishop, M. J. 1974. A preliminary report on the Middle Pleistocene mammal bearing deposit 

of Westbury-sub-Mendip, Somerset. Proc. speleol. Soc, Bristol, 13 : 301-318. 

1975- Earliest record of Man's presence in Britain. Nature, Lond. 253 : 95-97. 

Blackmore, H. p. & Alston, E. R. 1874. On Fossil Arvicolidae. Proc. zool. Soc Lond. 

1874 : 460-471, 3 figs. 


Blore, J. D. 1966. The excavation of Lynx Cave, 1962- 1964. A preliminary report. 

Newsl. Peakland archaeol. Soc, Stockport, 21 : 17-19, 35. 
BoYLAN, P. J. 1967. The Pleistocene mammalia of the Sewerby-Hessle buried cliff. East 

Yorkshire. Proc. Yorks. geol. Soc, Leeds, 36 : 115- 125. 
Bramwell, D. i960. The vertebrate fauna of Dowel Cave - final report. Newsl. Peakland 

archaeol. Soc, Stockport, 17 : [9-12, 23] (pages unnumbered). 
1964. The excavations at Elder Bush Cave, Wetton, Staffs. North Staffs. J. Fid Studs, 

Keele, 4 : 46-60. 
1970. The tenth report on Fox Hole Cave, High Wheeldon. Bull. Peakland archaeol. 

Soc, Stockport, 25 : 8-10. 
Brewster, T. C. M. 1963. The Excavation of Staple Howe. Malton, E. Riding Arch. Res. 

Brown, J. 1852. On the Upper Tertiaries at Copford, Essex. Q. Jlgeol. Soc Lond. 8 : 184- 

BuRCHELL, J. P. T. 1935. Evidence of a further glacial episode within the valley of the 
Lower Thames. Geol. Mag., London, 72 : 90-91. 

* Burleigh, R. 1972. Carbon 14 dating with application to dating of remains from caves. 

Stud. Speleol., London, 2 : 176-190. 
Campbell, J. B. & Sampson, C. G. 1971. A new analysis of Kent's Cavern, Devonshire, 

England. Univ. Ore. anthrop. Pap., Eugene, Oregon, 3 : 1-40. 
Carreck, J. N. 1957. A late Pleistocene rodent and amphibian fauna from Levaton, 

near Newton Abbot, South Devon. Proc Geol. Ass., London, 68 : 304-308. 
1959. In : Kerney, M. P. An interglacial tufa near Hitchin, Herts. Proc. Geol. 

Ass., London, 70 : 322-337. 
1966. Microtine remains from the Norwich Crag (Lower Pleistocene) of Easton Bavents, 

Suffolk. Proc Geol. Ass., London, 77 : 491-496. 
Catt, J. a. & Penny, L. F. 1966. The Pleistocene deposits of Holderness, East Yorkshire. 

Proc Yorks. geol. Soc, Leeds, 35 : 375-420. 
Chaline, J. 1972. Les Rongeurs du Pleistocene Moyen et Superieur de France. 410 pp., 

17 pis. Paris. 
Cheadle, R. W. 1876. On mammalian bones recently found at Crayford and Erith. 

Proc W. Lond. scient. Ass. Fid Club 1 : 70-71. 
Coleman, J. C. 1965. The Caves of Ireland. 88 pp., 21 figs, 8 pis. Tralee. 

* CooPE, G. R. 1974. Interglacial Coleoptera from Bobbitshole, Ipswich, Suffolk. /. geol. 

Soc. Lond. 130 : 333-340. 

* 1975. An ecological study of a temperate interlude in the middle of the Last Glaciation, 

based on Coleoptera from Isleworth, Middlesex. /. anim. Ecol., London. 44 : 365-391. 

, Shotton, F. W. & Strachan, I. 1961. A late Pleistocene fauna and flora from Upton 

Warren, Worcestershire. Phil. Trans. R. Soc, London, B 244 : 379-421. 

* Corbet, G. B. 1961. Origin of the British insular races of small mammals and of the 

'Lusitanian' fauna. Nature, Lond. 191 : 1037- 1040. 

* 1966. The Terrestrial Mammals of Western Europe. xi + 264pp. London. 

* , Cummins, J., Hedges, S. R. & Krzanowski, W. 1970. The taxonomic status of 

British water voles, genus Arvicola. J. Zool., Lond. 161 : 301-316. 
Cranbrook, Lord 1955a. Fossil Voles. Trans. Suffolk Nat. Soc, Ipswich, 9 : 48-49, 

3 figs [published anonymously]. 
1955b. Records of Mimomys. Trans. Suffolk Nat. Soc, Ipswich, 9 : 58 [published 

Davies, J.A. 1921. Aveline's Hole, Burrington Combe. Proc speleol. Soc, Bristol, I: 61-68. 
Dawkins, W. B. & Sanford, W. A. 1866-72. British Pleistocene Mammalia, 1 (Felidae). 

1+ 194 pp., 26 pis. Palaeontogr. Soc (Monogr.), London. 

* Dienske, H. 1969. Notes on difference between some external and skull characters of 

Microtus arvalis (Pallas, 1778) and of Microtus agrestis (Linnaeus, 1761) from the Nether- 
lands. Zool. Meded. Leiden 44 (6) : 83-108. 


* Ellerman, J. R. & Morrison-Scott, T. C. S. 1966. Checklist of Palaearctic and Indian 

Mammals. 2nd ed. 810 pp., map. London, British Museum (Natural History). 

* Evans, P. 1971. Towards a Pleistocene time-scale. In : The Phanerozoic Time-scale - a 

Supplement. Spec. Pubis geol. Soc. Lond. 5 : 123-351. 
Evans, W. 1907. Mammals of the Edinburgh or Forth area. Proc. R. phys. Soc. Edinb. 

16 : 387-405- 
1913. Lemming remains in south-east Scotland. Scott. Nat., Edinburgh, 17 : 97-100, 

I fig. 

* Fahlbusch, v. 1969. Pliozane und Pleistozane Cricetinae (Rodentia, Mammalia) aus 

Polen. Acta zool. cracov. 14 (5) : 99-137, pis 8-18. 
Falconer, H. See Murchison, C. 1868. 

* Franks, J. W. i960. Interglacial deposits at Trafalgar Square, London. New Phytol., 

Oxford, 59 : 145-152. 

Eraser, F. C. & King, J. E. 1954. Faunal remains. Ch. 3 in : Clark, J. G. D. Excava- 
tions at Star Carr : 70-95. Cambridge. 

Friant, M. 1962. Sur le Castor plicidens F. Maj. de I'epoque des Palafittes. Jb. naturh. 
Mus. Bern 1960-62 : 117- 125, pis 1-2. 

* Giebel, C. G. a. 1847. Fauna der Vorwelt mit steter Beriicksichtigung der lebenden Thiere 1 

(i). Die Saugethiere. 467 pp. Leipzig. 

* Gromov, I. M. 1965. In : Gromov, I. M., Bibikov, D. I., Kalabukhov, N. I. & Meier, 

M. N. Nezemnye belichi (Marmotinae) . Fauna SSSR, Moscow & Leningrad, 3 (2) : 1-466. 
HiNTON, M. A. C. 1900a. On the Pleistocene deposits of the Ilford and Wanstead district, 

Essex. Essex Nat., Buckhurst Hill, 11 : 161 -165. 
1900b. The Pleistocene deposits of the Ilford and Wanstead district. Proc. Geol. Ass., 

London, 16 : 271-281. 

1901. Excursion to Grays Thurrock. Proc. Geol. Ass., London, 17 : 141 -144. 

1907a. On the existence of the alpine vole {Microtus nivalis Martins) in Britain during 

Pleistocene times. Proc. Geol. Ass., London, 20 : 39-58. 
1907b. Note on the occurrence of the alpine vole {Microtus nivalis) in the Clevedon 

Cave deposit. Proc. Bristol Nat. Soc. (4) 1 : 190- 191. 
1910a. Some new late Pleistocene voles and lemmings. Ann. Mag. nat. Hist., London, 

(8) 6 : 34-39. 
1910b. A preliminary account of the British fossil voles and lemmings ; with some 

remarks on the Pleistocene climate and geography. Proc. Geol. Ass., London, 21 : 489-507. 

1 9 1 2 . Note on the lemming remains from the Arctic Bed at Angel Road . In : Warren, 

S. H. A Late Glacial stage in the valley of the River Lea. Q. Jl geol. Soc. Lond. 68 : 249. 

1914. On some remains of rodents from the Red Crag of Suffolk. Ann. Mag. nat. Hist., 

London, (8) 13 : 186-195, i pi. 
1915. Note on British fossil species of Apodemus. Ann. Mag. nat. Hist., London, (8) 

15 : 580-584. 

1920. Description of the rodent teeth. In : Trechmann, C. T. On a deposit of inter- 
glacial loess and some transported pre-glacial clays on the Durham Coast. Q. J I geol. Soc. 
Lond. 75 : 201. 

1921. Note on the remains of small mammals obtained from Aveline's Hole, Burrington 

Combe, Somerset. Proc. speleol. Soc, Bristol, 1 : 74-78. 
1923a. Diagnoses of species of Pitymys and Microtus occurring in the Upper Freshwater 

Bed of West Runton, Norfolk. Ann. Mag. nat. Hist., London, (9) 12 : 541-542. 

1923b. Note on the rodent remains from Clacton-on-Sea. Q. Jl geol. Soc. Lond. 79 : 626. 

1924. Notes on the rodent remains from Aveline's Hole. Proc. speleol. Soc, Bristol, 

1925. Preliminary note upon the mammalian remains from Merlin's Cave. Proc. 

speleol. Soc, Bristol, 2 : 156-158. 
1926a. The Pleistocene mammalia of the British Isles and their bearing upon the date 

of the glacial period. Proc. Yorks. geol. Soc, Leeds, (n.s.) 20 : 325-348. 


HiNTON, M. A. C. 1926b. Monograph of the Voles & Lemmings [Microtinae) Living and 
Extinct. 1 : xvi + 488 pp., 15 pis, no figs. London, British Museum (Natural History). 

1952. In : Allison, J., Godwin, H. & Warren, S.H. Late-glacial deposits at Nazeing 

in the Lea Valley, North London. Phil. Trans. R. Soc, London, B 236 : 169-240. 

, Kennard, a. S. & Newton, E. T. 1900. Contributions to the Pleistocene geology of 

the Thames Valley. I. The Grays Thurrock area, part i. Essex Nat., Buckhurst Hill, 

11 : 336-370- 
& White, G. 1902. Note on the occurrence of Microtus intermedins in the Pleistocene 

deposits of the Thames Valley. Proc. Geol. Ass., London, 17 : 414-415. 
Jackson, J. W. 1909. On the discovery of the remains of lemmings in Dog Holes, Warton 

Crag. Lanes. Nat., Darwen, 2 : 227-229. 
1910. On the vertebrate fauna found in the cave-earth at Dog Holes, Warton Crag. 

Lanes. Nat., Darwen, 2 : 323-332. 
191 2. Additional notes on the Pleistocene fauna of Dog Holes, Warton Crag. Lanes. 

Nat., Darwen, 4 : 420-422. 

1929a. Remains of lemmings in Derbyshire caves. Naturalist, Hull 1929 : 105-107. 

1929b. Report on the animal remains found in Kilgreany Cave, Co. Waterford. Proc. 

speleol. Soc., Bristol, 3: 137-152. 
1932. [Dicrostonyx in Welsh caves.] Proc. Dyserth Distr. Fid Club, Llanfairfechan, 

54 [not seen]. 
1934. The rodent remains from the Pin Hole Cave. Rep. Br. Ass. Advmt Sci., Aberdeen : 


1947- The Upper Pleistocene fauna and its relation to the Ice Age. Proc. Lpool geol. 

Soc. 19: 165-183. 

* Janossy, D. 1954. Fossile Microtinen aus dem Karpathenbecken. I. Lemminge. Annls 

hist.-nat. Mus. natn. hung., Budapest, n.s. 5 : 39-48, i pi. 
* 1 96 1. Die Entwicklung der Kleinsaugerfauna im Pleistocan (Insectivora, Rodentia, 

Lagomorpha). Z. Sdugetierk., Berlin, 26 : i-ii. 
* 1962. Vorlaufige Mitteilung iiber die Mittelpleistozane Vertebratenfauna der Tarko- 

Felsnische (NO-Ungarn, Biikk-Gebirge) . Annls hist.-nat. Mus. natn. hung., Budapest, 

54: 155-176. 
Johnson, J. P. & White, G. 1900. Some new sections in, and contributions to, the fauna of 

the river drift of the Uphall estate, Ilford, Essex. Essex Nat., Buckhurst Hill, 11 : 157-160. 

* Kaup, J. J. 1839. Description d'Ossements Fossiles de Mammiferes {&c.) 5: gi-iig. Darmstadt. 
Kennard, A. S. 1944. The Crayford brickearths. Proc. Geol. Ass., London, 55 : 121 -169. 
1945-46. The early digs in Kent's Hole, Torquay, and Mrs Cazalet. Proc. Geol. Ass., 

London, 56 : 156-213. 
Kerney, M. p. 1971. Interglacial deposits in Barnfield Pit, Swanscombe, and their 
molluscan fauna. /. geol. Soc, London, 127 : 69-93. 

* King, W. B. R. & Oakley, K. P. 1936. The Pleistocene succession in the lower parts of 

the Thames Valley. Proc. prehist. Soc, Cambridge, (n.s.) 2 : 52-76. 

* KoENiGSWALD, W. voN 1970. Mittclplcistozane Kleinsaugerfauna aus der Spaltenfiillung 

Petersbuch bei Eichstatt. Mitt, bayer. St. Paldont. Hist. Geol., Munich, 10 : 407-432. 
1972. Sudmer-Berg-2, eine Fauna des friihen Mittelpleistozans aus dem Harz. Neues 

Jb. Geol. Paldont. Abh., Stuttgart, 141 : 194-221. 
1973- Veranderungen in der Kleinsaugerfauna von Mitteleuropa zwischen Cromer und 

Eem (Pleistocan). Eiszeitalter Gegenw., Ohringen, 23-24 : 159-167. 

* KoRMOS, T. 1934. Neue Insectenfresser, Fledermause und Nager aus dem Oberpliocan 

der Villdnyer Gegend. Foldt. Kdzl., Budapest, 64 : 296-321. 
KowALSKi, K. 1966. Stratigraphic importance of rodents in the studies on European 

Quaternary. Folia quatern., Krakow, 22 : 1-16. 
1967. Lagurus lagurus (Pallas 1773) and Cricetus cricetus (Linnaeus 1758) (Rodentia, 

Mammalia) in the Pleistocene of England. Acta zool. cracov. 12 : 111-122, 24 figs. 
* 1970. Variation and speciation in fossil voles. Symp. zool. Soc Lond. 26 : 149-160. 


Kretzoi, M. 1958. New names for Arvicolid homonyms. Annls Jiist.-nat. Mus. natn. 

hung., Budapest, 50 : 55-58. 
1965. Die Nager und Lagomorphen von Voigstedt in Thiiringen und ihre chronologische 

Aussage. Paldont. Abh. Berl. A 11 : 585-660, 2 pis, 2 figs. 

* KuRTEN, B. 1969. Pleistocene Mammals of Europe. viii + 3i7pp. London. 
Lamplugh, G. W. 1891. Final report of the Committee . . . appointed for the purpose of 

investigating an ancient sea-beach near Bridlington Quay. Rep. Br. Ass. Advmt Sci., 
Leeds : 375-377- 
Lankester, E. R. 1864. On new mammalia from the Red Crag. Ann. Mag. nat. Hist., 
London, 14 : 353-360, pi. 8. 

* Laugel, a. 1862. La faune de Saint- Prest pres Chartres (Eure et Loire). Bull. Soc. g&ol. 

Fr., Paris, (2) 19 : 709-718, 3 figs. 

* Lehmann, U. 1953. Eine Villafranchiano-Fauna von der Erpfingen Hohle (Schwab. Alb). 

Neues Jb. Geol. Paldont. Mh., Stuttgart, B 1953 : 437-464. 

* 1957- Weitere Fossilfunde aus dem altesten Pleistocan der Erpfingen Hohle (Schwa- 

bische Alb.). Mitt. geol. Stinst. Hamb. 26 : 60-99. 

Lydekker, R. 1885-87. Catalogue of the Fossil Mammalia in the British Museum {Natural 
History). 1 (Primates, Chiroptera, Insectivora, Carnivora and Rodentia), 1885, xxx + 
268 pp. 5 (Tillodontia, Sirenia, Cetacea, Edentata, Marsupiaha, Monotremata and Supple- 
ment), 1887, xxxv + 345 pp. London. 

Lyell, C. 1840. On the Boulder Formation or drift and associated freshwater deposits 
composing the mud cliffs of eastern Norfolk. Lond. Edinb. phil. Mag., London, (3) 16 : 345- 
380, 18 figs. 

Major, C. I. F. 1902. [Exhibition of, and remarks upon, some jaws and teeth of Pliocene 
voles (Mimomys, gen. nov.).] Proc. zool. Soc. Lond. 1902 (i) : 102-107. 

1908a. The mammalian fauna of the Forest Bed. Geol. Mag., London, (5) 5 : 329-330. 

1908b. On species of Castor, fossil and recent. Proc. zool. Soc. Lond. 1908 : 630-635. 

* Mandach, E. V. 1938. Skelettreste von Dicrostonyx groenlandicus Trail als Inhalt von 

Raubvogelgewollen. Meddr Grenland, Copenhagen, 112 (4) : 1-54, 22 pis. 

* Matthews, L. H. 1952. British Mammals, xii-f 410 pp., 16 + 48 pis. London. 

* Mehely, L. v. 1914. Fibrinae Hungariae. Die ternaren und quartaren Wurzelzahnigen 

Wiihlmause Ungarns. Annls hist.-nat. Mus. natn. hung., Budapest, 12 : 155-243. 

* Miller, G. S. 1912. Catalogue of the Mammals of Western Europe. xv+ 1019 pp. London, 

British Museum. 
Mitchell, G. F. 1969. Pleistocene mammals in Ireland. Bull. Mammal Soc. Br. Isl., 

Birmingham, 31 : 21-25. 
* , Penny, L. F., Shotton, F. W. & West, R. G. 1973. A correlation of Quaternary 

deposits in the British Isles. Spec. Rep. geol. Soc. Lond. 4 : 99 pp. 
Montagu, I. 1924. On the remains of Fen Beaver in the Sedgwick Museum. Proc. zool. 

Soc. Lond. 1924 : 1081-1086. 
MuLLiNS, E. H. 1913. The ossiferous cave at Langwith. /. Derbysh. archaeol. nat. Hist. 

Soc, Derby, 35 : 137-153, 6 pis. 
MuRCHisoN, C. (ed.) 1868. Palaeontological memoirs and notes of the late Hugh Falconer. 

1 : lvi + 590 pp., 34 pis. 2 : xiii + 675 pp., 38 pis. London. 
Nehring, a. 1875. Fossile Lemminge und Arvicolen aus den Diluviallehm von Thiede 

bei Wolfenbiittel. Z. ges. Naturw., Berlin, 45 : 1-28, pi. I. 
Newton, E. T. 1881. Notes on the Vertebrata of the pre-glacial Forest Bed Series of the 

East of England. Geol. Mag., London, (2) 8 : 256-259. 
1882a. The Vertebrata of the Forest Bed Series of Norfolk and Suffolk. viii + 143 pp., 

19 pis. Mem. geol. Surv. U.K., London. 
1882b. On the occurrence of Spermophilus beneath the glacial till of Norfolk. Geol. 

Mag., London, (n.s. 2) 9 : 51-54, pi. 2. 
1890a. On the occurrence of lemmings and other rodents in the brick-earth of the 

Thames Valley. Geol. Mag., London, (n.s. 3) 7 : 452-455, 8 figs. 


Newton, E. T. 1890b. On some new mammals from the Red and Norwich Crags. Q. Jl 
geol. Soc. Lond. 46 : 444-453, pi. 18. 

1891. The Vertebrata of the PHocene deposits of Britain. xi+ 137 pp., 10 pis. Mem. 

geol. Surv. U.K., London. 

1892. On a skull of Trogontherium cuvieri from the Forest Bed of East Runton, near 

Cromer. Trans, zool. Soc. Lond. 13 : 165-175, pi. 19. 
1894. The vertebrate fauna collected by Mr Lewis Abbot from the fissure near Ightham, 

Kent. Q. J I geol. Soc. Lond. 50 : 188-210, pis 10-12. 

1899a. [Exhibition of remains of Mus Lewisi from Ightham.] Proc. zool. Soc. Lond. 

1899 : 381. 

1899b. Additional notes on the vertebrate fauna of the rock-fissure at Ightham (Kent). 

0. Jl geol. Soc. Lond. 55 : 419-429, pi. 28. 

1902. Trogontherium from a Pleistocene deposit in the Thames Valley. Geol. Mag., 

London, (n.s. 4) 9 : 385-388. 
— — 1909. Hamster remains from the Norfolk Forest Bed. Geol. Mag., London, (5) 

6 : 110-113. 

1916. Trogontherium from the Pleistocene of Copford, Essex. Geol. Mag., London, (6) 

3 : 322-323, 4 figs. 

* Norton, P. E. P. 1967. Marine Mollusca in the Early Pleistocene of Sidestrand and the 

Royal Society borehole at Ludham, Norfolk. Phil. Trans. R. Soc, London, B 253 : 161-200. 

* Oken, L. 1815-16. Okens Lehrbuch der Naturgeschichte. Ill Zoology 1: xxviii + 842 + 

xviii + iv pp., 40 pis. 2 : xvi+ 1270+ (2) pp., i tab. ; pp. 843-50 (suppl. to 1). 
OsBORN, H. F. 1922. Pliocene (Tertiary) and early Pleistocene (Quaternary) mammaUa of 
East Anglia, Great Britain, in relation to the appearance of Man. Geol. Mag., London, 

* OvEY, C. D. (ed.) 1964. The Swanscombe Skull. Occ. Pap. R. anthrop. Inst., London, 

20 : xii-j-215 pp., 25 pis. 
Owen, R. 1846. A History of British Fossil Mammals, and Birds, xlvi-l-560 pp., 236 figs. 

1869. On the distinction between Castor and Trogontherium. Geol. Mag., London, 

6 : 49-56, pi. 3. 4 figs- 
Palmer, L. S. 1934. Some Pleistocene breccias near the Severn Estuary. Proc. Geol. Ass., 

London, 45 : 145 -161. 
Pasquier, L. 1972. Etude d'une population de Mimomys savini Hinton, 1910 (Arvicolidae, 

Rodentia) provenant de I'Upper Freshwater Bed (Quaternaire Ancien d' Angle terra). 

Mammalia, Paris, 36 : 214-225. 
Peach, B. N. & Horne, J. 1917. The bone cave in the valley of Allt nan Uamh (Burn of 

the Caves) near Inchnadamph, Assynt, Scotland. Proc. Roy. Soc. Edinb. 37 : 327-349. 
Pernetta, J. C. 1966. Report on the findings at Etches' Cave, Dowel Dale, Derbyshire. 

Newsl. Peakland archaeol. Soc, Stockport, 21 : 11 -16, 35. 
& Handford, p. T. 1970. Mammalian and avian remains from possible Bronze Age 

deposits on Nornour, Isles of Scilly. /. Zool., Lond. 162 : 534-540. 

* Pike, K. & Godwin, H. 1952. The interglacial at Clacton-on-Sea. Q. Jl geol. Soc. Lond. 

108 : 261-272. 

* PoMEL, A. 1848. Description de la tete du Castoroides Ohioensis Forster par Wyman. 

Archs Sci. phys. nat., Geneva, 9 : 165-167. 

* Reichstein, H. 1963. Beitrag zur systematischen Gliederung des Genus Arvicola Lacepede 

1799. Z. Zool. Syst. EvolForsch., Frankfurt a.M., 1 : 155-204. 
Reid, C. 1890. The Pliocene deposits of Britain, viii-l-326 pp., 5 pis, 48 figs. Mem. geol. 
Surv. U.K., London. 

* RzEBiK, B. 1968. Crocidura Wagler and other Insectivora (Mammalia) from the Quaternary 

deposits of Tornewton Cave in England. Acta zool. cracov. 13 : 251-263. 
Sanford, W. a. 1870a. On the rodentia of the Somerset caves. Proc. Somerset archaeol. 
nat. Hist. Soc, Taunton, 1868-69 : 51-57, i pi- 


Sanford, W. a. 1870b. On the rodentia of the Somerset caves. Q. Jl geol. Soc. Lond. 26 : 

Savage, R. J. G. 1963. Martin AHster Campbell Hinton, 1883- 1961. Biogr. Mem. Fellows 

R. Soc, London, 9 : 155-170, i pi. 

1966. Irish Pleistocene mammals. Ir. Nat. J., Belfast, 15 : 117- 130. 

ScHARFF, R. F. 1906. The exploration of the caves of County Clare. Trans. R. Ir. Acad., 

Dublin, B 33: 1-76. 
, Coffey, G., Cole, G. A. J. & Ussher, R. J. 1903. The exploration of the Caves of 

Kesh, County SHgo. Trans. R. Ir. Acad., Dubhn, B 32 : 171-214, pis 9-11. 
, Seymour, H. J. & Newton, E. T. 1918. The exploration of Castlepook Cave, County 

Cork. Proc. R. Ir. Acad., Dublin, B 34 : 33-72 + map. 

* ScHAUB, S. 1930. Quartare und jungtertiare Hamster. Abh. schweiz paldont. Ges., Basel, 

49 (6) : 1-49, 2 pis. 
ScHLOSSER, M. 1902. Beitrage zur Kenntniss der Saugetierreste aus den siiddeutschen 

Bohnerzen. Geol. palaeont. Abh., Berlin, 9 : 117-258, 5 pis. 
ScHREUDER, A. 1929. Conodontes {Trogontherium) and Castor from the Tegelen Clay, 

compared with Castoridae from other localities. Archs Mus. Teyler, Haarlem, (3) 6 : 99- 

320, 16 pis, 16 figs. 

1931- Conodontes, Trogontherium and other Castoridae. Palaeont. Z., Berlin, 13 : 148- 

176, 6 figs. 

1950. Microtinae from the Middle Gravels of Swanscombe. Ann. Mag. nat. Hist., 

London, (12) 3 : 629-635. 
195 1. The three species of Trogontherium with a remark on Anchitheriomys . Archs 

neevl. Zool., Leiden, 8 : 400-433, pis 20-23. 

* SiDOROWicz, J. 1964. Comparison of the morphology of representatives of the genus Lemmus 

Link, 1795, from Alaska and the Palaearctic. Acta theriol., Bialowieza, 8 : 217-226. 
Singer, R., Wymer, J., Gladfelter, B. G. & Wolff, R. G. 1973. Excavation of the 
Clactonian industry at the golf course, Clacton-on-Sea, Essex. Proc. prehist. Soc, Cam- 
bridge, 39 : 6-74. 
♦Southern, H. N. (ed.) 1964. The Handbook of British Mammals, xxi + 465 pp., 60 pis. 

* Sparks, B. W. & West, R. G. 1972. The Ice Age in Britain, xvii + 302 pp., 44 pis, figs. 

Spencer, H. E. P. 1956. The Hoxne mammalian remains. In : West, R. G. The 

Quaternary deposits at Hoxne, Suffolk. Phil. Trans. R. Soc, London, B 239 : 265-365. 
1964. The contemporary mammalian fossils of the Crags. Trans. Suffolk Nat. Soc, 

Ipswich, 12 : 333-344- 
1966. New mammalian fossils from the Red Crag. Trans. Suffolk Nat. Soc, Ipswich, 

13: 154-156. 
Stevens, E. T. 1869. Drift series ~ mammalian remains. In : Some Account of the 

Blackmore Museum 2 : 109- 115. Devizes. 
Stores, C. 1904. Palaeolithic implements from the Shelly Gravel Pit at Swanscombe, 

Kent. Rep. Br. Ass. Advmt Sci., Southport : 803-804. 
Stringer, C. B. 1975. A preliminary report on new excavations at Bacon Hole Cave. 

Gower, Swansea, 26 : 32-37. 
Stuart, A. J. 1974. Pleistocene history of the British vertebrate fauna. Biol. Rev., 

Cambridge, 49 : 225-266. 

* Sutcliffe, a. J. i960. Joint Mitnor Cave, Buckfastleigh. Trans. Torquay nat. Hist. Soc. 

13 (I) : 1-26. 

1964. The mammalian fauna. In : Ovey, C. D. (ed.), The Swanscombe Skull. Occ. 

Pap. R. anthrop. Inst., London, 20 : 85-1 11. 

& BowEN, D. Q. 1973. Prehminary report on excavations in Minchin Hole, April- 
May 1973. Newsl. Pengelly Cave Stud. Trust 21 : 12-25. 


SuTCLiFFE, A. J. & CoLLiNGS, H. D. 1972. Gnawed bones from the Crag and Forest 
Bed of East Anglia. Suffolk nat. Hist., Ipswich, 15 : 497-498. 

& Zeuner, F. E. 1962. Excavations in the Torbryan Caves, Devonshire. I. 

Tornewton Cave. Proc. Devon archaeol. Explor. Soc, Torquay, 5 : 127-145, pis 26-27. 

* Turner, C. 1970. The Middle Pleistocene deposits at Marks Tey, Essex. Phil. Trans. R. 

Soc, London, B 257 : 373-440. 
UssHER, R. J. 1910. Cave hunting. Ir. Nat., DubUn, 19 : 37-43. 
, Seymour, H. J., Newton, E. T. & Scharff, R. F. 1908. On the Cave of Castlepook, 

near Doneraile, Co. Cork. Rep. Br. Ass. Advmt Sci., Dubhn : 697 ; Geol. Mag., London (5) 

5 : 462-463 ; Ir. Nat., Dublin, 17 : 231-232. 

* ViRET, J. 1954. Le loess a bancs durcis de Saint- Valher. Nouv. Archs Mus. Hist. nat. 

Lyon 4 : 1-200, 33 pis. 
Warren, S. H. 1916. Further observations on the Late Glacial, or Ponders End, stage of 

the Lea Valley. Q. Jl geol. Soc. Lond. 71 : 164-182. 

1955- The Clacton (Essex) Channel deposits. Q. Jl geol. Soc. Lond. Ill : 283-307. 

♦West, R. G. 1961. The Glacial and Interglacial deposits of Norfolk. Trans. Norfolk 

Norwich Nat. Soc. 19 : 365-375. 

1968. Pleistocene Geology and Biology (&c.). xiii + 377 pp., 16 pis, figs. London. 

* 1969. Pollen analyses from interglacial deposits at Aveley and Grays, Essex. Proc. 

Geol. Ass., London, 80 : 271-282. 
* 1972. Land -sea-level changes during the Pleistocene. In : A discussion on problems 

associated with the subsidence of southeastern England. Phil. Trans. R. Soc, London, 

A 272 : 87-98. 
* , Lambert, A. & Sparks, R. W. 1964. Interglacial deposits at Ilford, Essex. Phil. 

Trans. R. Soc, London, B 247 : 185-212. 

* & Wilson, D. G. 1966. Cromer Forest Bed Series. Nature, Lond. 209 : 497-498. 

Whitaker, W. 1889. The geology of London and of part of the Thames Valley 1. ix-t- 

556 pp. Mem. geol. Surv. U.K., London. 

* Wymer, J. 1957. A Clactonian flint industry at Little Thurrock, Grays, Essex. Proc. 

Geol. Ass., London, 68 : 159-177. 

1962. Excavations at the Maglemosian sites at Thatcham, Berkshire, England. Proc. 

prehist. Soc, Cambridge, 28 : 329-361. 
* 1974. Clactonian and Acheulean industries in Britain - their chronology and signifi- 
cance. Proc. Geol. Ass., London, 85 : 391-421. 

Zeuner, F. E. 1945-59. The Pleistocene Period, its Climate, Chronology and Faunal Suc- 
cessions. 1st edition (1945), 322 pp. London (Ray Society). 2nd edition (1959), 447 pp., 
80 figs. London. 
* 1954. Riss or Wiirm? Eiszeitalter Gegenw., Ohringen, 4-5 : 98-105. 

Zimmermann, K. 1959. Ober eine Kreuzung von Unterarten der Feldmaus Micratus 
arvalis. Zool. Jb., Jena, (Abt. Syst.) 87 : 1-12. 


The page numbers of the principal references are printed in bold type. An asterisk (*) denotes 
a figure. Tab. 12 faces p. 122. 

Acheulean industry 50-1 alluvial deposits 41, 53, 74, tab. 12 ; see 

Africa 82 localities 

alder 52 Alopex see fox (arctic) 

Alice and Gwendoline Caves 68 ; see Eden- Alston, E. R. 37 

vale Alveston Fissure 45, 67, 93, no, tab. 12, 129 

Allocricetus 66, 86-7, 127 amphibia 59 

cf. bursae 63-4, 66, 86-7, loi, tab. 12, Angel Road 45, 59, 88, tab. 12 

125-6 Anghan 41-2, 47-8, 76, 106, 123 

Allophaiomys pliocaenicus 105-7, ^^2 Antian 41, 43 



Apodemus 81-4 
flavicollis 62, 67, 70-1, 83-4, tab. 12, 129 
lewisi 83-4 
sylvaticus 48, 52-3, 59, 61, 63-72, 81-3, 

tab. 12, 122-7, 129 
whitei 52, 81-3 
sp. 69, 81, 83 
Arctic Freshwater Bed 48 
Arvicola 37-9, 52-3, 64, 69, 99-102, 103-5 I 
see Mimomys-A rvicola lineage 
faunas 61, loo-i, tab. 12, 124, 127 
abbotti 103 
agrestis 109 
amphibia 37, 102-3 
ampJiibius 95, 98, 102-3 

terrestris 103 
arvalis 37, 106 
arvaloides 106 

bactonensis 48, 100, 102, 104 
cantiana 48, 51-3, 57, 60-2, 69, 72, 100*, 

loi, 102-3, 104-5, tab. 12, 122-5 
cantiana-terrestris transition form 60, 64-5, 

67, 69, 72, 100*, IOI-2, 103, 125-7 
cantiana /terrestris complex 104-5 
glareolus 92 

greeni 48, 62, 100, 102, 104-5 
gregalis 107, 118 
gulielmi 37, 87 
intermedia 37, 95, 98-9 
praeceptor 53, 100, 102, 104 
pratensis 92 
ratticeps 117 
riparia 92 
sapidus 99 
terrestris 37, 59, 62, 64-5, 69-71, 99, 100*, 

101-2, 103-4, 105, tab. 12, 127-9 
sp. 54, 63-7, 69, 72, 125 
Asia 76, 80, 82, 84-5, 87, 89, 94, 100, 105, 

112, 118, 120-1 
Asia Minor 78 
Astian 41, 43, 78, 81 
Austria 106, 109, 118 
Aveley 45, 49-50. 53- 55-7. 69, 93, 101-2, 

tab. 12, 125-6, 130 
Aveline's Hole 45, 70-1, 81, 83-4, 88, 90, 93, 

104, no, 112, 117-8, tab. 12 
Avon, Warwickshire 72 
Azov Sea 79 

Bacon Hole 45, 67, 81, 93, 104, no, 117 
Bacton 45, 46*, 48, 78-9, 104, tab. 12 
Baker's Hole 49-50, 59-60 
cold stage 127 

Ballymote 45 

Ballynamintra Cave 45, 68, 82, 88, tab. 12 

bank vole see Clethrionomys 

Banwell Cave 45, 66, tab. 12, 126 

Barnfield Pit 51-3, 80, 90, 92-3, 101-2, tab. 

12, 123-4, 130 ; see Swanscombe 
Barntick Cave 68, 82 ; see Edenvale 
Barrington 45, 69, 72, 101-3, no, tab. 12, 

Bath 112 
Baventian 41, 43 
Beaker Age 71 
bear 60, 62-3, 65 
brown 63-4, 68 
Bear Stratum (Tornewton Cave) 63-4, 66 
beaver 37, 39, 81 ; see Castor, Trogontherium 
Beccles 45 

Beckford 45, 72, 113, tab. 12, 128-30 
Beestonian 41, 47-8, tab. 12 
Belgium 83, 96 
Bering Strait 92, 120 
Berkshire 45 ; see Thatcham 
Berwickshire 45 ; see Middlestots Bog 
Betfia, Romania 95 
Bhukhureea, Mrs M. N. M. 130 
Biedermann'scher Stbr. loi 
birch 56 

birds of prey 60, 69 ; see owls 
Bishop, M. J. 108, 129 
bison 56 

Black Mould (Kent's Cavern) 62 
Blackmore, H. P. 37 
Bleadon Cave 45, 70-1, 76-7, no, 117, 119, 

tab. 12 
Bobbitshole 45, 69, 72, 104, no, tab. 12, 127 
Bos see ox, giant 
bovids 62-3 

Bramerton 43, 45, 46*, 95, 98, tab. 12 
Bramwell, D. 130 
Brandon 89 
Brassington 45 

Brean Down 45, 72, 88, no, tab. 12 
Breccia (Kent's Cavern) 62, tab. 12 
brickearth 48, 50, 53-4, 56-8 
Briggs, D. J. 72, n3 
Bristol Museum 107 
Bristol University Spelaeological Society 81, 

88, 91 
British Museum (Natural History) 36, 58, 

60-1, 66, 72, 97-8, 102, 107, 113, 116, 

Brixham Cave 45, 70-1, 93, 104, no, 112-3, 

119, tab. 12 
Bronze Age 128 ; see Nornour 



Buckfastleigh 45 

Buckinghamshire 45 ; see Marlow 
Burchell, J. P. T. 60 
Burrington 45 
Burwell 45 
Buxton 45 

Calcareous Group (Westbury) 60-1, 122, 124 
Cambridge Fens 45, 72, 80, tab. 12 
Cambridge, P. 96 (footnote) 
Cambridgeshire 45 ; see locahties 
Canada 92 ; see North America 
Canis see wolf 

lupus mosbachensis 61 
Cappagh 45 
carbon-14 (i*C) dated sites 36, 58, 68-9, 72, 

113, tab. 12, 127-9 
Carnforth 45 
Carrant Brook 72 
Carreck, J. N. 50, 54, 56, 59-60, 69, 89, 91, 

93, 104, no, 127, 129 
Castlepook Cave 45, 68-9, 82, 88, 91, tab. 12, 

Castletownroche Cave 45, 68-9, 88, 91, 

tab. 12 
Castor 39, 54, 69, 79-81 

europaeus 79 

fiber 37, 43, 47-8, 52-3, 61-2, 70-3, 79-81, 
tab. 12, 122-8 
plicidens 81 

plicidens 79-81 

trogontherium 78 

veterior 79-80 

sp. 53. 79 
Castoridae 78-81 
Catacombs 68 ; see Edenvale 
Caucasus 83, 115 

cave deposits 41-2, 56, 58, 60-9, tab. 12 ; 
see localities 

disturbed 67 
Cave Earth (Kent's Cavern) 62, loi, 104, 

tab. 12 
Cervus see red deer 
Chaline, J. 39 
channels, buried 48-9 
Cheddar 45 
China 82, 87 
Chinese Turkestan 121 
Chudleigh (Fissure) 45, 70-1, 88, 90, 93, 104, 

no, 117, 119, tab. 12 
Citellus 75, 128 ; see Spermophilus 

erythrogenoides 75 

erythrogenys 75 

eversmanni 75 

nogaici 75 

{Colobotis) superciliosus 75, 77 

(Urocitellus) polonicus 75-6 

primigenius 75 
sp. 76 
Clacton-on-sea 45, 49*, 52-3, 78-80, 93, 

101-2, 109, tab. 12, 123 
Clactonian industry 50-1, 53 
Clare, Co. 45 ; see Edenvale Caves 
Clarke, Dr A. S. 130 
classification 73-121 
'clay pebbles' 47 
Clethrionomys 92-4 

glareolus 37, 48, 52-3, 59-67, 70-2, 92-4, 

tab. 12, 122-7, 129 

nageri 94 
sp. 53. 67, 92 
Clevedon Cave 45, 66, 93, in, 115-7, tab. 12, 

Coelodonta see rhinoceros, woolly 
Coffey Cave see Keshcorran 
Colchester 45 

Colobotis see Spermopfiilus superciliosus 
Connaberry 45 
Conodontes Boisvilletti 78 
coombe rock 50, 59-60 ; see solifiuxion 

Coope, G. R. 72, 113, 127-8, 130 
Copford 45, 69, 78-9, tab. 12, 123 
Coralline Crag 41, 96 (footnote) 
Corbet, Dr G. B. 58, 72, 93, iio-i, 113, 115, 

Corbicula Bed 54, 89, 92, 125 
Cork, Co. 45 ; see localities 
Corstorphine 45, 73, 88, no, tab. 12 
Covehithe 43, 45, 46*, 96, tab. 12 
Cow Cave 45, 66, 115, tab. 12, 126 
Cowside Cave No. 3 45, 70-1, 104, no, 118, 

tab. 12 
Cranbrook, Earl of 98 
Crags 37, 41-2, 60 ; see Coralline, Icenian, 

Norwich, Red, Shelly and Weybourne 

Crayford 38, 45, 49*, 50, 54, 60, 64, 66, 75, 

92, 101-3, in, 115, 117-8, tab. 12, 

Gravel 50 
Creag nan Uamh Cave 45, 68, 88, no, 118, 

tab. 12 
Cresswell 45 ; see Pin Hole Cave 
Cricetidae 85-121 
Cricetulus migratorius 87 
Cricetus 48, 85-6, 127 



cricetus 48, 63-6, 85-6, tab. 12, 122, 124-6 
major 86 
runtonensis 86 
runtonensis 86 
songarus 37, 86-7 
vulgaris Runtonensis 37, 85 
Crocidura 64, 126 
Crocuta see hyaena 
Cromer 45, 46*, 78, 98, tab. 12 
Cromer Forest Bed Series (Norfolk) 36-7, 
40-1, 43, 46-8, 74, 77, 79, 85, 94, 97-8, 
105, tab. 12 
Cromerian stage 40-1, 47, 60-2, 69, 78, 82-3, 
85, 93, 96, 98, loi, 105-8, 114, 116, 
tab. 12 
sensu stricto 47-8, 52, 61-2, 73, tab. 12, 
Currant, A. P. 130 
Cyrnaonyx 64, 126 

Czechoslovakia 76, 86-7, 89, 96-9, 106, 109, 
114, 116 

Dama see fallow deer 

deep-sea cores 40 

deer see fallow, giant, red deer, reindeer, &c. 

Denbighshire 45 ; see Lynx Cave 

Denmark 113 

Derbyshire 45 ; see localities 

Devensian 41-2, 115, tab. 12, 127-8 

Devon 45, 126 ; see localities 

Diabroticus Schmerlingi 78 

Dicerorhinus see rhinoceros, narrow-nosed 

etruscus 61, 69 

hemitoechus 56 

kirchbergensis 56 
Dicrostonyx 67, 69, 72-3, 87-90, 92, iii, 
122-3, 127-9 

gulielmi 87, 90 

henseli 87, 90 

hudsonius 90 

simplicior 89 

torquatus 37, 54, 59, 62-8, 70-2, 87-90, 
91-2, tab. 12, 125-6, 128 

sp. 61, 87, 89, 124 
Dierden's Pit 51 ; see Swanscombe 
'Diluvium' (Tornewton Cave) 62-4, 66, no, 

Dipoides Lydekkeri 78 
Diss 45 ; see Hoxne 
distribution 73-121 
Dnepr, river 121 
Dog Holes Cave 45, 70-1, 77, 81, 88, 90, 93, 

104, 109-10, 112-3, 117, 119, tab. 12 
domestic animals 84 

Domnitz 42 

Doneraile 45 

dormouse see Muscardinus, Glis 

Dove Holes 60 

Dowel Cave 45, 70-1, 74, 82-3, 88, 91, 93, 

117, 119, tab. 12, 128 
Dowel Dale 45 ; see Dowel Cave, Etches' 

Drepanosorex savini loi 
Droitwich 45 

Earl Sterndale 45 

East Anglia 42, 46*, 98 ; see localities 

deposits in 42-8, 69, 72 
East Runton 38, 45, 46*, 47-8, 78-80, 96, 

98, tab. 12, 122 
East Wickham 115 

Easton Bavents 43, 45, 46*, 95, 98, tab. 12 
Ebbsfieet 49-50, 59-60 
Eburonian 41, 43 

Edenvale Caves 45, 68, 82, 88, tab. 12 
Edinburgh 45 ; see Corstorphine 
Edmonton 48, 49*, 59 
Edrom Parish 45 ; see Middlestots Bog 
Eemian 40-2, 100 
Elder Bush Cave 45, 70-1, 88, 91, 93, 104, 

no, 117, tab. 12 
elephant, straight-tusked 56-7 
Elsterian 41, 61, 100 
Ennis 45 
Eocene 43 
Eppelsheim 76 
Equus see horse 

mosbachensis 61 
Erith 38, 45, 49*, 50, 54, 75, 87, 89-90, 92, 

115, 117, tab. 12, 125 
Erkenbrechtsweiler loi 
Erpfingen i and 2 loi 
Essex 45 ; see localities 
Etches' Cave 45, 70-1, 83, 88, 91, 117, tab. 

12, 128 
Europe, continent of 36, 39, 43, 54, 74, 76-8, 
80, 82-5, 87, 89, 92, 94, 97, 99, 105-8, 
112-4, 116, 118, 120-2, 124, 126-8 
Evans, P., half-cycles 40, 42, 127 
Evotomys glareolus 92 

harrisoni 92, 94 

intermedius 95, 97-8 

kennardi 92, 94 

nageri 94 

sp. 92, 94 

Falconer, H. 37 
fallow deer 56, 63 


Felis see lynx 

gomhaszoegensis 6i 
Felixstowe 45, 46*, 78 
field-vole see Arvicola 
Finland 112 
fir cones, gnawed 74 

Fisherton 45, 72, 76, 88, 115, 117, tab. 12 
Fitch, Mr 96 (footnote) 
Flandrian 41, 59, tab. 12 
Flint bed 47 

Flintshire 45 ; see Gwaenysgor Cave 
Floodplain Terrace complex (Thames) 54, 

footprints 51 

Forest Bed 42, 46*, 48, 53 ; see Cromer 
Forest of Dean 115 
fox, arctic 72 

Fox Hole Cave 45, 70-1, 88, tab. 12, 128 
France 74, 76, 78-9, 83, 87, 89, 95-9, 105, 

109, 118, 120-1, 127 

Geldeston 45, 46*, 98 

geological background 40-2 

Geological Society of London 40, 50, 55 

Georgia 106, 116 

Germany 76, 79, 87, 92, 96-9, 106, 109, 114, 

116, 118, 124 
giant beaver see Trogontherium 
giant deer 68 

Gippingian, Gipping cold stage 41, 56 
glacial till 48 

Glamorganshire 45 ; see Penard 
Gliridae, glirids 43, 77-8 
Glis glis 129 

Gloucestershire see Alveston Fissure 
glutton 63, 126 
Glutton Stratum (Tornewton Cave) 63-6, 

82, 85-6, 88-90, 92-3, 101-3, IIO-I, 

115, 117, 120, tab. 12, 125-6 
cold stage tab. 12, 125, 127 
Gough's Cave 45, 66, 104, no, 115, 117, 

tab. 12, 126 
Grays Thurrock 38, 45, 49*, 50, 53-4, 57, 69, 

80-1, 93, 101-3, 109, III, 115, tab. 12, 

Great Doward Cave 45, 70-1, 77, 81, 88, 91, 

93, no, 117, 119, tab. 12 
Green, Rev. C. 48 
Greenland 92 
Gromov, I. M. 75, 77 
ground squirrels see Spermophilus 
Gulo see glutton, wolverine 
Giinz, Giinz-Mindel 76 
Gwaenysgor Cave 45, 70-1, 81, 88, 93, tab. 12 

Hackney (Marshes) 45, 48, 49*, 59, 88, tab. 12 
hamster see Cricetus, Allocricetus 
Happaway Cave 45, 70-1, 82-3, 93, 104, no, 

119, tab. 12 
Happisburgh 45, 46*, 47-8, tab. 12 
Harborough Cave 45, 70-1, 91, tab. 12 
Harkstead 45, 69, 72, 102-3, no. n?. tab. 

12, 125 
harvest mouse see Micromys 
Hay Wood Rockshelter 45, 70-1, 80, 118, 

tab. 12 
Helmsley 45 
Heppenloch loi 

herbivores, large, trampling by 54 
Herefordshire 45 ; see localities 
Hertfordshire 45 ; see localities 
Hessle 45, 69, 102, 104, tab. 12 
High Wheeldon Hill 45 ; see Fox Hole Cave 
hillwash 49 ; see Northfleet 
Hinton, M. A. C. 35-40 

rodent faunas 40-1 
hippopotamus. Hippopotamus 50, 56-9, 63, 

67, 69, 86, 93, 102, 121, 125-7 
history of rodent faunas 121 -9 
Hitchin 45, 69, 81, 93, tab. 12 
Holland 43, 79, 83, 96-8, 116, 118 
Holocene 41, 59-60, 62-4, 67, 70-3, 77, 80, 

82-3, 93, 100, 102, 104, 113, tab. 12, 

Holsteinian Interglacial 40-2, 79, 83, 95, 105, 

Homo, hominids 51 ; see industries, man 
Homotherium 62 

latidens 61 
Hooker, J. J. 130 
hornbeam 56 
horse 56, 63, 72 
Hoxne 42, 45, 69, 78-9, 81, 90, 92, 104, tab. 

12, 123, 130 
Hoxnian Interglacial 40-2, 52-3, 55-6, 65, 

69, 78-80, 82, 92-3, loo-i, 104-8, no, 

116, tab. 12, 123, 125 
Hull see Kingston-upon-Hull 
Hunas loi 
Hundsheim 61, loi 
Hungary 74, 76, 87, 89-90, 92, 95-8, 106, 

109, 114, 116, 118, 120 
Huntspill Cut 45, 73, 118, tab. 12 ; see 

Hutton Cave 45, 58, 66-7, 86-7, 90, 93, tab. 

12, 125-6 
hyaena 62-3, 67, 69 
Hyaena Stratum (Tornewton Cave) 63-6, 

104, no, 121, tab. 12, 125-6 


Hypudaeus bucklandii 109 

spelaeus 103 
Hystrix 43, 122 

Iberian peninsula 99 ; see Spain 

ice covering Britain 113 

ice wedges 58 

Icenian Crag 41, 43, tab. 12, 122 

Ightham Fissures 37-8, 41, 45, 67, 76-7, 81, 

83-4, 88, 91, 93-4, loi, 104, no, 112-3, 

117, 119, tab. 12 
Ilford 45, 49-50. 53. 55-^> 60, 69, 80, 103, 

109, tab. 12, 125-7 
Inchnadamph 45, 67 ; see Creag nan Uamh 

Ingress Vale 51-2, 79, 81, 93, 100-2, tab. 12, 

123-4 '< s^^ Swanscombe 
insects 36, 58, 72, 113, 127-8 
Institute of Geological Sciences Museum 104 
Ipswich 45, 72 

Ipswich Museum 72, 97, 103-4, no. ^i? 
'Ipswichian' 57, 63, 67, 69, 72, tab. 12, 125-7 
Ipswichian stage (interglacial) 40-2, 50, 

53-7, 82, 91, 124-5, 127 
Ireland 42, 91, 129 ; see counties 
caves 68-9 
rodents in 121 
voles in in -2 
Iron Age 84 ; see Staple Howe 
Isleworth 45, 48, 49*, 58, 117, 119, tab. 12, 

Italy 95, 97-8, 106, 114 ; see Val d'Arno 

Jewell, Dr J. 130 

Joint Mitnor Cave 45, 56, 67, 82, 93, 101-3, 
iio-i, tab. 12, 126 
interglacial tab. 12, 125-6 

Kadzielnia 98 

Kalkberg, Nagyharsanyberg 76 

Kent 45, 50 ; see localities 

Kent's Cavern 45, 60, 61-2, 80, 83, 88, 91, 93, 

loi, 104, 107, no, 117, 119, tab. 12, 122 
Kerney, M. P. 51, 60, 63 
Keshcorran Caves 45, 68, 82, 89, tab. 12 
Kessingland 45, 46*, 78-9, 98, tab. 12 
Kilgreany Cave 45, 68-9, 82, 84, 89, in, 

tab. 12 
King Arthur's Cave 45, 70-1, 88, no, 117, 

119, tab. 12 
Kingston-upon-Hull 45 ; see Hessle 

Kinlock 45 

Kirkdale Cave 45, 56, 67, 81, 93, 104, no, 

tab. 12 
Koenigswald, W. von 36, 39, 64, 67, 69, loi, 

Mimomys-Arvicolai3im:ial groups of 100- 1, 

tab. 12, 124, 127 
Kongprusy loi 
Kopet-Dag 115 
Kyson 45, 96 

Lagurus 120-1, 127 

lagurus 63-6, 120-1, tab. 12, 125-6 
Lancashire 45 ; see Dog Holes Cave 
Langwith Cave 45, 70-1, 74, 76, 82, 84, 88, 

91, 93, 104, no, 112, 117, 119, tab. 12 
Last Glaciation 50, 54, 56, 58-60, 62-4, 

67-72, 76, 82, 89-94, 102, 104, III, 113, 

118-21, 126, 127-8, 129 
Last Interglacial 40, 42, 50, 54-5, 56*, 59, 

80, 89, 93, 102, 104 
last interglacial 65, 78 
Lea Valley 38, 41, 48-60, 49*, 58-9, 128-9 ; 

see Nazeing, Ponders End, Water Hall 

Farm, &c. 
Lebanon 115 
lemming 54, 64, 67, 122, 127-9 ; see Lemmus, 

Dicvostonyx, Lagurus 
Lemmus 37, 67-8, 90-2, 123-4, 127-9 

lemmus 54, 59, 62-6, 68, 70-1, 90-2, tab. 

12, 125-6, 128 
norvegicus 37, 90 
torquatus 87 

sp. 52, 61, 69, 90, 122-4 
Levallois industries 49-50, 54 
Levaton Cave 45, 67, 70-1, 82, 104, no, 112, 

117, 119, tab. 12, 128 
Limerick, Co. 45 ; see Red Cellar Cave 
lion 56, 63 
Little Thurrock 50 

localities with fossil rodents 42-73, 44-5* 
Loch of Marlee 45, 73, 80, tab. 12 
London 48, 58 ; see Hackney, Angel Road, 

Lough Gur 45 

Lower Crayford Brickearth 50 
Lower Loam (Swanscombe) 42, 52, 102, 108, 

116, 123-4, 130 
Lower Thames Valley province 48 
Lowestoftian 41, 123 
Ludhamian 41 
lynx 68 
Lynx Cave 45, 70-1, 82, 88, no, 118, tab. 12 



MacEnery, J. 62 

Macoma balthica 43 

Main Chamber see Tornewton Cave 

Major, C. I. Forsyth 75 

Male, H. C. 66 

mammals other than rodents 56, 60, 72 ; see 

mammoth 50, 56-60, 62, 67-9, 72, 87, 129 
Mammuthus 59 ; see mammoth 

primigenius 50 

trogontherii 50, 56 
man, introductions to Britain by 83-4, 129 

remains, human iii 
Marlow 45, 48, 58, 113, 117, tab. 12, 129 
mastodon 43 
Mauer 61, loi 
May hew, D. 130 
Mediterranean 115 

Medway, Upper Floodplain Terrace of 54 
Megaceros see giant deer 
melt- water deposits 59-60 
Menapian 41 

Mendips see Bleadon Cave, &c. 
Merlin's Cave ( = Wye Cave) 45, 70-1, 88, 91, 

93, loi, 104, no, 117, 119, tab. 12 
Mesolithic 62, 128 ; see Star Carr, Thatcham 
Micromys 66, 84 

minutus 65, 84, tab. 12, 129 

sylvaticus 81 
Microtinae 37-8, 43, 107, in 
Microtius {Pitymys) arvalidens 106 
Microtus 69, 105, 107-8, 108-20 

agrestis 37, 53-4, 58-9, 62-9, 72, 108, 
109-12, 113, 119, tab. 12, 125-7, 129 
neglectus 109 

of agrestis group 53 

agrestoides 109, in 

anglicus 60, 118-20, 128 

arvalinus 48, 52, 61, 108-9, iio-i, tab. 12, 

arvalis 58, 60, 72, 108-9, 112-3, tab. 12, 

sp. {arvalis I agrestis group) 52, 60, 70-1, 
108, 109, III 

corneri 109, 112 -3 

eversmanni 120 

glareolus 92 

gregalis 58-60, 62-4, 66, 70-1, 107, 113, 
118-20, tab. 12, 126-8 

intermedins 102 

malei 114-6, 118 

nivalinus 114 

nivalis 37, 54, 59-60, 63-6, 72, 114-6, 
tab. 12, 125-7 

nivaloides 48, 114, tab. 12, 122-4 
oeconomus 37, 54, 58-9, 62-3, 65-8, 70-3, 
108, 113, 115-6, 117-8, tab. 12, 125-8 
orcadensis 112 -3 
ratticepoides 48, 52, 116-7, 118, tab. 12, 

ratticeps 68, 116-7 
subnivalis 114 
tianschanicus 120 
sp. 66, 69 

{Pitymys) arvalidens 105-6 
sp. 106 
Middle East 100 
Middle Terrace of Thames interglacial tab. 

12, 125, 127 
Middlesex 45 ; see Isleworth, Angel Road, 

Ponders End 
Middlestots Bog 45, 73, 80, tab. 12 
Midlothian 45 ; see Corstorphine 
Mimomys 60, 95-100 

cantiana, cantianus 100, 102 
intermedins 98-9 
majori 98-9 
milleri 98-9 

newtoni 43, 47, 97-8, tab. 12 
petenyi 97 

pliocaenicus 43, 47, 95-6, tab. 12, 122 
polonicus 96 
reidi 43, 97, tab. 12, 122 
savini 47-8, 52, 69, 98-9, 100- 1, 105, tab. 
12, 122-4 
fauna tab. 12, 124 ; see Arvicola 

sp- 43 
Mimomys-Arvicola lineage 36, 100, tab. 12, 

Minchin Hole 45, 67, 103, no, tab. 12,127, 129 
Mindel 76 
Miocene 43 

Mitchell, Professor G. F. 130 
mollusca 43, 51-2, 54, 56, 58, 63, 125 
Monastirian, Late 40, 42, 50, 55 

Main 42, 50, 55 
Mongolia 121 

'monkey gravel' 48, 73, 122 
Mosbach 61, loi, 122 
multituberculate origin of rodents 38 
Mundesley 45, 46*, 48, 75-6, 78-9, tab. 12, 

Arctic Bed tab. 12, 123 
Muridae, murids 43, 81-4 
Murston 45, 88, tab. 12 
Mus 84 

Abhotti 83 

Lewisi 83 



musculiis 8i, 84, tab. 12, 129 

songarus 86 

sylvaticus 81 

sp. 81 
Muscardinus 48, 77-8 

avellanarius 67, 70-1, 77-8, tab. 12, 129 
Myodes lemmus 90 

torquatus 87 

narrow-skulled vole see Microtus gregalis 
Nazeing 45, 48, 49*, 59, 82, 88-9, 91, 93, 104, 

118-9, tab. 12, 128 
Neolithic 62, 83 
Netherlands see Holland 
Newhall Caves 68 ; see Edenvale 
Newton, E. T. 37 

Norfolk 38, 42, 45-6, 113 ; see localities 
Nornour 45, 73, 118, tab. 12 
North America 89-90, 92, 94, 118 
Northfleet 45, 49*, 50, 59-60, 93, 103, no, 

tab. 12, 126 
Norwich Crag 38, 41, 43, 46*, 47, 78, 95-8 ; 

see Icenian Crag 
Norwich Museum 96 (footnote) 

oak 56 

Orkney Islands 109, 112 -3 
Orkney vole 112 ; see Microtus arvalis 
Orsett Road 50, 53 

Ostend (Norfolk) 45, 46*, 48, 53, 74, 101-2, 
104, tab. 12, 123 
Forest Bed 47-8, ^2» 74-5- 9^, 122-3 
otter, clawless 64, 126 ; see Cyvnaonyx 
Otter Stratum (Tornewton Cave) 36, 64-5, 

101-3, tab. 12, 126 
Overstrand 45, 46*, 78, tab. 12 
Owen, R. 37 
owls 69, 112 
ox, giant 56 
Oxfordshire 45 ; see Sugworth 

palaeobotany 39-40, 46-8, 55, 57, 59, 124-7 ! 

see pollen, plants 
Palaeolithic 62-3, 66, 71 
Palaeoloxodon antiquus 59 ; see elephant, 

Palestine 105 
Panthera see lion 
Parapodemus coron. loi 
Paston 45, 46*, 78, tab. 12 

Pastonian (Forest Bed) 41, 43, 47-8, 78-9, 

96, 98, tab. 12, 122 
Peak District 36 

Peakland Archaeological Society 67, 128, 130 
Penard 45 
Penkridge 89 

Penultimate Glaciation 42 
'penultimate' Glaciation 54, 56, 59, 64-5, 

112, 114, 121, 125 
Perpignan 78 
Persia 105 

Perthshire 45 ; see Loch of Marlee 
Petersbuch loi 
Phodopus 87 

sanfordi 86 

songorus 86-7 
Picken's Hole 45, 70-1, 76, 117, 119, tab. 12, 

Pin Hole Cave 45, 70-1, 76-7, 82-3, 88, 91, 

93, 104, no, 117, 119, tab. 12 
pine 52, 56 

pine vole see Pitymys 
Pitymys 105-8 

arvalidens 105 

arvaloides 48, 52, 105, 106, 107, tab. 12, 

gregaloides 48, 61-2, 105-6, 107, tab. 12, 
sp. 69 
plants 53, 58, 122, 127 ; see palaeobotany 
Pleistocene 67, 74, 82-5, 87, 91-2, 112, 
121-9, tab. 12 
generaHzed British sequence 41 

Lower 43, 60, 98, 121 

lower middle 80, 87, 89, 92, 94-5 

Upper 83 
Pliocene 41, 43, 112, 122 ; 'Pliocene' 37, 40 
Pliomys 95, 123 

episcopalis 61, 95, loi, tab. 12, 122, 124 

lenki loi 
Plumpstead 115 

Plunkett Cave 68 ; see Keshcorran 
Poland 76, 87, 89, 92, 96-8, 106, 116 
pollen 36, 52, 55-6, 89, 128-9 ; see palaeo- 

zonation of 'Ipswichian' sites 56*, 57-8 
Ponders End 45, 48, 49*, 50, 59, 88, tab. 12 
post-Ipswichian 67 

post-Pleistocene 68, 73 ; see Holocene 
'preglacial' deposits 37 
pre-'Ipswichian' 65-7 
Prestatyn 45 
Pfezletice 10 1 
Pyrenees 115 



radiocarbon dating see carbon- 14 
Rangifev see reindeer 
rat see Rattus 
Rattiis 84 

norvegicus 84, tab. 12, 129 

rattus 67, 70-1, 74, 84, tab. 12, 129 
Red Cellar Cave 45, 68, 89, tab. 12 
Red Crag 41, 43, 46*, 78-80, 95, tab. 12, 122 

basal nodule bed 43 
red deer 56, 63, 67 
reindeer 58, 63, 67-9, 72, 126 
Reindeer Stratum (Tornewton Cave) 63-6, 
88-9, 93, loi, 104, no, 117, 119, tab. 12 
Reuverian 41 
rhinoceros 56, 59, 63 

narrow-nosed 63 

woolly 62, 67, 69, 72 
Riss 42, 76, 100, 112, 121 
Rodent Earth (Westbury) 60-1, 122, 124 
rodent localities on British mainland tab. 12 
Romania 92, 95-6, 109, 114, 116 
Romano-British 71 
root vole see Microtus oeconomus 
Rowberrow Cavern 45, 70-1, 82, 88, 93, 

tab. 12 
Russia 76, 87, 95-8, 106, 116 

Saale 42, 50 

Saalian 41 

sabre-toothed cat 62 ; see Homotherium 

Saint-Prest, Chartres 79 

St Vallier, France 81 

Salisbury 45 

Salwarpe river 72 

Sanford, W. A. 37 

Savin collection 97 

Scandinavia 92, 112-3 ; see Sweden, &c. 

Scarborough 45 

Scarcliffe 45 ; see Langwith Cave 

Scilly Isles 45, 73, 118, tab. 12, 128 ; see 

Sciuridae 73-7 

Sciurus 73-4 ; see squirrel 

vulgaris 67, 70-1, 74, tab. 12, 129 
whitei 48, 73-4, tab. 12, 122, 124 
hungaricus 74 

Scotland 73, 80, 128 ; see localities 
caves in 67-8, 118 

Sealy, D. L. F. 130 

Selsey 45, 69, 80, tab. 12 

Settle 45 

Seven Kings Station 55-6, 58 

Severn river 72 

Shell Cottages, Thorpe, Aldringham 96 (foot- 
Shelly Crag (East Runton) 38, 41, 47 
shrew, white-toothed 64-5, 126 ; see Croci- 

Siberia 96-8, 105 
Sieveking, G. de G. 60 
Siliceous Group (Westbury) 60-1, 124 
silver fir 56 
Simons, J. 58, 66, 130 
Singer, Dr R. 130 

Sizewell 43, 45, 46*, 78, 95, 97, tab. 12 
Sligo, Co. 45 ; see Keshcorran Caves 
snow vole see Microtus nivalis 
solifluxion deposits 59-60 
Somerset 45 ; see localities 

caves 37 

Levels see Huntspill Cut 
Sorex runtonensis loi 

savini 10 1 
Soviet Union see Russia 
Spain 87 

Spencer, H. E. P. 96 (footnote), 130 
Spermophilus 48, 54, 75, 76 

altaicus 75-6 

citillus 76 

eversmanni 75-6 

erythrogenoides 37, 75-7 

erythvogenys 75 

sp. 75, 128 

(Colobotis) superciliosus 70-2, 75, 76-7, 
tab. 12, 127 

{Urocitellus) parryi 76 
primigenius 54, 75-6, 77, tab. 12, 125-7 
undulatus 76 
squirrels 48, 74 ; see Sciurus 
Staffordshire 45 ; see Elder Bush Cave 
Stalagmite (Kent's Cavern) 62 
Staple Howe 45, 72, 80, tab. 12 
Star Carr 45, 73, 80, tab. 12, 128 
Steinheim an der Murr 124 
Stenocranius 120 ; see Microtus gregalis 
steppe lemming see Lemmus lemmus 
Stoke Tunnel Beds 45, 72, 103, tab. 12 
Stuart, A. J. 39, 60, 69, 72, 81, 93, 102-4, 

no, 117, 129 
studies, history of 37-40 
Stutton 45, 69, 72, 81, 101-3, no, tab. 12, 

Sudmer Berg 2 loi 
Suffolk 42-3, 45 ; see localities 
Sugworth 45, 69, 98, tab. 12 
Siissenborn loi 
Sussex 45 ; see West Wittering, Selsey 



Sutcliffe, Mrs A. J. 130 

Sutherland 45 ; see Creag nan Uamh Cave 

Sutton 43, 45, 46*, 79, tab. 12 

Swaffley 45 

Swanscombe 36, 38-9, 42, 45, 49*, 50, 51-2, 
53. 59, 78-81, 83, 90, 92-3, 100-2, 
108-11, 115-6, tab. 12, 123-4 

Swanton Morley 45, 69, 72, 81, 93, 101-3, 
tab. 12, 126 

Sweden 78, 83 

Switzerland 87, 89, 118, 120 

Symond's Yat 45 

Talpa europaea loi 

minor loi 
Taplow terrace 55 
Tarko 61, 74, loi 
terrace deposits 41-2, 48-60, tab. 12 ; see 

Thames 48-60, 49*, 62 

High Terrace 38, 41, 49, 51-3 ; see 

Lower 51-8 

Middle Terraces 36, 38, 41, 49, 55, 57, 125 
early 53 
interglacial see Middle Terrace of 

Thames interglacial 
late 54, 125 
Middle Terrace/Floodplain Terrace prob- 
lem 55-8 
Third Terrace 38, 41, 54, 55 
Upper 58 

Upper Floodplain Terrace 55-8 
Thatcham 45, 73, 80, 104, tab. 12 
Thorpe, Aldeburgh, Suffolk 96 (footnote) 
Thorpe, Norfolk 43, 45, 46*, 78, 96 (incl. 

footnote), tab. 12 
Thorpeness 96 (footnote) 
Thurnian 41, 43 

Tiglian 41, 43, 78, 83, 96-8, 122 
tooth-enamel 81, 100 
Torbryan 45 

Tornewton Cave 36, 45, 62-5, 66-7, 81, 
84-90, 92-3, 101-4, iio-i, 115, 117, 
119-21, tab. 12, 125-6 
Torquay 45 

Trafalgar Square 55-8, 127 
Treacher collection 58, 113 
Trimingham 43, 45, 46*, 97, tab. 12 
Trogontherium boisvilletti 43, 47-8, 52-3, 69, 
78-9, tab. 12, 122-4 
cuvieri 37, 53, 78-9 

minus 37, 43, 78, 122 
sp. 78 
Turner, C. 54 

Ukraine 74, 76, 114 

Uphall Estate 109 

Uphill Cave 45, 70-1, 91, 119, tab. 12 

Upnor 45, 49*, 54, 80, tab. 12 

Upper Freshwater Bed (West Runton) 38, 

41, 48, 52, 73, 81, 83, 85, 93, 97-101, 

105, 107-8, 114, 116, 124 
Upper Langwith 45 
Upper Rodent Stratum (Tornewton Cave) 

Uppony 10 1 
Upton Warren 45, 69, 72, 88, tab. 12, 128 

interstadial complex 128 
Urals 83 
Urocitellus 125 ; see Spermophilus primi- 

Ursus see bear 

deningeri 61 

Val d'Arno, Italy 80-1, 96 

Vejlby I and II mild stages (Denmark) 40, 42 

Villafranchian 41, 43, 47, 76, 81, 92, 95-6, 

Viscount Melville Sound 92 
Voigtstedt loi 

voles 38-9, 47, 51, 69 ; see Arvicola, 
Mimomys, Microtus 
in Ireland ? 69, 129 

Waalian 41, 43 

Waechter, J. d'A. 51, 130 

Wales, south 113 

Waltonian 41 

Warthe 42 

Warton Crag 45 ; see Dog Holes Cave 

Water Hall Farm (Pit) 45, 48, 49*, 58-9, no, 

115, 117, tab. 12, 126, 130 
Waterford, Co. 45 ; see localities 
water-rat, water-vole see Arvicola 
Weichsel 42 
Weichselian 41 
West, R. G. 47, 130 
West Runton 38, 41, 45, 46*, 47-8, 52, 60-1, 

73, 78-9, 81, 83, 85, 93, 97-100, 105-8, 

114, 116, tab. 12, 122-4 
West Thurrock 50 



West Wittering 45, 93, tab. 12 

Westbury-sub-Mendip (fissure) 36, 45, 60-1, 
80-1, 87, 89-90, 92-3, 95, 101-2, 105, 
107-8, 121, tab. 12, 122-4, 129 

Westbury stage 108, tab. 12, 122, 123 

Wetton 45 

Weybourne Crag 38, 41, 43, 46*, 47, 95-7 ; 
see Icenian Crag 

Whitchurch 45 

White Sea 120 

Whitechurch 45 

Willment's Gravel Pit 58, 128, 130 

Wilson, Mrs G. 130 

Wiltshire 45 ; see Fisherton 

Wintringham 45 

wolf 62-3 

Wolff, R. G. 69, 81, 90, 104, 130 

Wolstonian stage 40-2, 50, 52, 54-5, 124-5 

wolverine 126 ; see glutton 

wood mouse see Apodemus sylvaticus 

Woodbridge 43, 45, 46*, 78-9, tab. 12 

Worcestershire 45 ; see localities 

Wretton 91 

Wiirm 42, 76, 121 

Wye Cave see Merlin's Cave 

Xenocyon lycaonoides 61 

yellow-necked mouse see Apodemus flavi- 

Yorkshire 45 ; see localities 
Yugoslavia 98, 106, 109, 114 


Department of Palaeontology 
British Museum (Natural History) 
Cromwell Road 
London SW7 5BD 


Institute of Systematic and 

Experimental Zoology 
Polish Academy of Sciences 

Accepted for publication 28 October 1975 





1. Cox, L. R. Jurassic Bivalvia and Gastropoda from Tanganyika and Kenya. 
Pp. 213 ; 30 Plates ; 2 Text-figures. 1965. out of print. 

2. El-Naggar, Z. R. Stratigraphy and Planktonic Foraminifera of the Upper 
Cretaceous — Lower Tertiary Succession in the Esna-Idfu Region, Nile Valley, 
Egypt, U.A.R. Pp. 291 ; 23 Plates ; 18 Text-figures. 1966. ^^ii. 

3. Davey, R. J., DowNiE, C, Sarjeant, W. a. S. & Williams, G. L. Studies on 
Mesozoic and Cainozoic Dinoflagellate Cysts. Pp. 248 ; 28 Plates ; 64 Text- 
figures. 1966. £8.20. 

3. Appendix. Davey, R. J., Downie, C, Sarjeant, W. A. S. & Williams, G. L. 
Appendix to Studies on Mesozoic and Cainozoic Dinoflagellate Cysts. Pp. 24. 
1969. 95p. 

4. Elx^iott, G. F. Permian to Palaeocene Calcareous Algae (Dasycladaceae) of 
the Middle East. Pp. iii ; 24 Plates ; 16 Text-figures. 1968. out of print. 

5. Rhodes, F. H. T., Austin, R. L. & Druce, E. C. British Avonian (Carboni- 
ferous) Conodont faunas, and their value in local and continental correlation. 
Pp- 313 \ 31 Plates ; 92 Text-figures. 1969. £13.10. 

6. Childs, a. Upper Jurassic RhynchoneUid Brachiopods from Northwestern 
Europe. Pp. 119 ; 12 Plates ; 40 Text-figures. 1969. £5.25. 

7. Goody, P. C. The relationships of certain Upper Cretaceous Teleosts with 
special reference to the Myctophoids. Pp. 255 ; 102 Text-figures. 1969. 


8. Owen, H. G. Middle Albian Stratigraphy in the Anglo-Paris Basin. Pp. 164 ; 
3 Plates ; 52 Text-figures. 1971. £7.20. 

9. Siddiqui, Q. a. Early Tertiary Ostracoda of the family Trachyleberididae 
from West Pakistan. Pp. 98 ; 42 Plates ; 7 Text-figures. 1971. £9.60. 

10. Forey, p. L. a revision of the elopiform fishes, fossil and Recent. Pp. 222 ; 
92 Text-figures. 1973. £ii.35- 

11. Williams, A. Ordovician Brachiopoda from the Shelve District, Shropshire. 
Pp. 163 ; 28 Plates ; 11 Text-figures ; no Tables. 1974. £12.80. 

Printed in Great Britain hyjohn Wright and Sons Ltd. at The Stonehridge Press, Bristol BS4 5NU 




2 4 SEP 1976 






GEOLOGY Vol. 27 No. 3 

LONDON: 1976 











British Museum (Natural History) 

Pp. 149-268 ; 4 Plates 



GEOLOGY Vol. 27 No. 3 

LONDON: 1976 


(natural history), instituted in 1949, is 
issued in five series corresponding to the Scientific 
Departments of the Museum, and an Historical series. 

Parts will appear at irregular intervals as they 
become ready. Volumes will contain about three or 
four hundred pages, and will not necessarily be 
completed within one calendar year. 

In 1965 a separate supplementary series of longer 
papers was instituted, numbered serially for each 

This paper is Vol. 27, No. 3, of the Geological 
iPalaeontological) series. The abbreviated titles of 
periodicals cited follow those of the World List of 
Scientific Periodicals. 

World List abbreviation : 
Bull. Br. Mus. nat. Hist. (Geol.) 

ISSN 0007-1471 

Trustees of the British Museum (Natural History), 1976 


Issued 23 September, 1976 Price £8.80 


By THE LATE F. E. ZEUNER and the late F. J. MANNING 


Synopsis ...... 

I. Introduction ..... 
II. Terminology and Classification . 

III. Material studied .... 

IV. Systematic Palaeontology: Synonomic list of Fossil Bees 

(excluding Apidae) .... 
Superfamily Apoidea Ashmead 
Family Colletidae Bingham 
Family Halictidae Ashmead 
Subfamily Halictinae Ashmead 
Genus Cyrtapis Cockerell . 

Cyrtapis anomalus Cockerell 
Genus Halictus Latreille 

Halictus ruissatelensis Timon-David 
Halictus florissantellus Cockerell 
Halictus miocenicus Cockerell 
Halictus scudderiellus Cockerell 
Halictus schemppi (Armbruster) 
Family Andrenidae Latreille 
Subfamily Andreninae Latreille 
Genus Andrena Fabricius . 
Andrena wrisleyi Salt . 
Andrena clavula Cockerell 
A ndrena grandipes Cockerell 
Andrena hypolitha Cockerell . 
Andrena lagopus Latreille 
A ndrena percontusa Cockerell 
Andrena sepulta Cockerell 
A ndrena primaeva Cockerell . 
Andrena spp. indet. 
Genus Lithandrena Cockerell 

Lithandrena saxorum Cockerell 
Genus Pelandrena Cockerell 

Pelandrena reducta Cockerell 
Subfamily Panurginae Leach . 
Genus Libellulapis Cockerell 

Libellulapis antiquorum Cockerell 
Libellulapis wilmattae Cockerell 
Family Andrenidae incertae sedis 

Andrenidae gen. et sp. indet. 
Family Melittidae Dumeril 

Subfamily Ctenoplectrinae Cockerell 
Genus Ctenoplectrella Cockerell . 
Ctenoplectrella dentata Salt . 
Ctenoplectrella viridiceps Cockerell 






Genus Glyp tapis Cockerell . 

Glyptapis fuscula Cockerell . 
Glyptapis mirabilis Cockerell 
Glyptapis neglecta Salt 
Glyptapis reducta Cockerell . 
Glyptapis reticulata Cockerell 
Subfamily Melittinae Dumeril 
Genus Melitta Kirby . 

Melitta willardi Cockerell 
Family Melittidae incertae sedis . 
Dasypoda (s.l.) sp. indet. 
Family Megachilidae Latreille . 
Subfamily Lithurginae Newman 
Genus Lithurge Latreille 

Lithurge adamitica (Heer) 
Subfamily Megachilinae Latreille . 
Tribe Anthidiini Michener . 
Genus Anthidium Fabricius 

Anthidium mortuum (Meunier) 
Anthidium exhumatum Cockerell 
Anthidium scudderi Cockerell 
Genus Dianthidium Cockerell 

Dianthidium tertiarium Cockerell 
Genus Lithanthidium Cockerell . 
Lithanthidium pertriste Cockerell 
Tribe Megachilini Latreille . 
Genus Heriades Spinola 

Heriades bowditchi Cockerell . 
Heriades halictinus Cockerell 
Heriades laminarum Cockerell 
Heriades mersatus Cockerell . 
Heriades mildredae Cockerell 
Heriades priscus Cockerell . 
Heriades saxosus Cockerell . 
Genus Megachile Latreille . 

Megachile amaguensis Cockerell 
Megachile praedicta Cockerell 
Megachile sp. indet. 
Genus Osmia Panzer . 

Osmia carbonum Heyden 
Osmia antiqua Heer 
Osmia nigra sp. nov. . 
Osmia sp. indet. . 
Family Megachilidae incertae sedis 
'Apiaria' dubia Germar 
Megachilidae gen. et sp. indet. 
Family Anthophoridae Dahlbom 
Subfamily Xylocopinae Latreille 
Tribe Ceratinini Latreille 
Genus Ceratina Latreille 

Ceratina disrupta Cockerell . 
Tribe Xylocopini Latreille . 
Genus Xylocopa Latreille . 





Xylocopa friesei Statz . 
Xylocopa hydrobiae Zeuner , 
Xylocopa jurinei (Heer) 
Xylocopa senilis Heer . 
Subfamily Anthophorinae Dahlbom 
Tribe EucERiNi Latreille 
Genus Tetralonia Spinola . 

Tetralonia berlandi Theobald 
Tribe Anthophorini Dahlbom 
Genus Anthophora Latreille 

Anthophora melfordi Cockerell 
Genus Anthophorites Heer . 

A nthophorites gaudryi Oustalet 
Anthophorites thoracica Heer 
Anthophorites longaeva Heer 
Anthophorites mellona Heer 
Anthophorites titania Heer 
Anthophorites tonsa Heer 
Anthophorites veterana Heer 
Tribe Melectini Westwood . 
Genus Promeleota Cockerell . 

Promelecta brevipennis Cockerell 
Family Anthophoridae incertae sedis 
'Anthophora' effossa Heyden. 
Anthophoridae gen. et sp. indet 
Superfamily Apoidea (except Apidae) incertae sedis 
Apoidea gen. et sp. indet. 
Fossils erroneously classified as Apoidea (excluding Apidae) 

Polistes kirbyanus Cockerell 
Larval Chambers .... 
'Genus' Celliforma Brown . 
Celliforma favosites Brown 
Celliforma spirifer Brown 
Celliforma germanica Brown 
Celliforma nuda Brown 
Celliforma bedfordi sp. nov. 
Celliforma septata sp. nov. 
Celliforma sp. indet. 
'Genus' Uruguay Roselli 

Uruguay auroranormai Roselli 
V. Systematic descriptions of fossil Apidae 
Family Apidae Latreille 
Chalcobombus group 

Genus Chalcobombus Cockerell 

Chalcobombus hirsutus Cockerell 
Chalcobombus humilis Cockerell 
Chalcobombus martialis Cockerell 
Tribe BoMBiNi Latreille 
Genus Bombus Latreille 

Bombus crassipes Novak 

Bombus abavus Heer . 

Bombus florissantensis (Cockerell) 





Bombus proavus Cockerell 
Tribe Meliponini Handlirsch 
Genus Sophrobombus Cockerell 

Sophrobombus fatalis Cockerell 
Genus Trigona J urine 

Subgenus Trigona Jurine 

Trigona {Trigona) erythra Schletterer 
Subgenus Hypotrigona Cockerell 

Trigona {Hypotrigona) gribodoi Magretti 
Subgenus Tetragona Lepeletier & Serville 
Trigona {Tetragona) succini (Tosi) 
Trigona {Tetragona) iridipennis Smith 
Subgenus indet. 

Trigona sicula (Tosi) . 
Tribe Apini Latreille 

Genus Electrapis Cockerell . 
Subgenus Electrapis Cockerell 

Electrapis {Electrapis) apoides Manning 
Electrapis {Electrapis) meliponoides (Buttel-Reepen) 
Electrapis {Electrapis) tornquisti Cockerell 
Subgenus Protobombus Cockerell 

Electrapis {Protobombus) indecisus (Cockerell) 
Electrapis {Protobombus) tristellus (Cockerell) 
Subgenus Roussyana Manning . 

Electrapis {Roussyana) palmnickenensis (Roussy) 
Electrapis {Roussyana) proava (Menge) 
Genus Apis Linnaeus 

Subgenus Synapis Cockerell 

Apis {Synapis) cuenoti Theobald . 
Apis {Synapis) henshawi Cockerell 

Apis {Synapis) henshawi dormiens subsp. nov. 
Apis {Synapis) henshawi henshawi Cockerell 
Apis {Synapis) henshawi kaschkei (Statz) 
Subgenus ^^25 Linnaeus. .... 

Apis {Apis) armbrusteri Zeuner 

Apis {Apis) armbrusteri armbrusteri Zeuner. 
Apis {Apis) armbrusteri scharmanni (Armbruster) 
Apis {Apis) armbrusteri scheeri (Armbruster) 
Apis {Apis) armbrusteri scheuthlei (Armbruster) 
Apis {Apis) melisuga (Handlirsch) 
Apis {Apis) mellifera Linnaeus 
Family Apidae incertae sedis 

'Apis' aquitaniensis de Rilly 
'Apis' dormitans Hey den 
'Apis' styriaca Pongracz 
'Bombus' carbonarius Menge 
'Bombus' muscorum Roussy 
'Bombus' pusillus Menge 
'Bombus' antiquus Heyden 
'Bombus' grandaevus Heer 
'Bombusoides' mengei Motschulsky 
Apidae gen. et sp. indet. 









Not Apidae . , . a.^.^ 

Lithohlatta lithopMla (Germar) 



Appendix, hy S. F. Morris .... 

Colletidae gen. et sp. indet. . 
Ctenoplectrella splendens Kelner-Pillault 
Anthiidini gen. et sp. indet. . 
Osmia sp. ..... 

Apoidea (? Anthophorinae) sp. 
Probombus hirsutus Piton (ms) 



Trigona {Hypotrigona) eocenica Kelner-Pillault 


Trigona {Hypotrigona) dominicana Wille & Chandler 256 

Trigona {Nogueirapis) silacea Wille 


Electrapis minuta Kelner-Pillault 


Apis catanensis Roussy 


? Apoidea sp. .... 


Not Apoidea .... 



Acknowledgements ..... 



References ....... 



Index . 



The status of 128 species of fossil bees and nests are reviewed, 19 of which are left under open 
nomenclature. Four species are removed from the Apoidea. A new species Osmia nigra 
sp. nov. and subspecies Apis {Synapis) henshawi dormiens subsp. nov. of bees, and two new 
'species' of bees' nests Celliforma bedfordi sp. nov. and Celliforma septata sp. nov., are described. 
An appendix covering the years 1960-74 has been added. 


hy S. F. Morris 

Until 1949, Professor Zeuner and Dr Manning had each been independently working 
on monographs of fossil bees, and until that time they were unaware of each other's 
work. But in 1949 they met and agreed to combine their efforts into a joint 
monograph. This collaboration continued until the death of Professor Zeuner in 
1963, followed by the death of Dr Manning in 1966. 

The major part of the present work was already finished by 1959, but the authors 
seemed to have had problems which prevented them completing it. The monograph 
lacks a projected part which was to have dealt with the evolution of the bees, but 
there are no extant manuscript notes for this. Certainly in 1964 Dr Manning was 
investigating a sphecid wasp from the Jurassic of Lerida Province, Spain, which he 
thought might be (or be closely related to) the ancestor of the bees. No written 
material of this work is extant either. 

Entries to the main body of the monograph ceased about 1959, so the editor has 
added an Appendix (p. 255) containing fossil bee references since that date, and a 
few earlier ones that the authors had missed which have since come to his notice. 
A few minor alterations to the body of the text have been made in order to bring it 
into line with modern taxonomic work. 


Although the monograph is intended to be a world-wide catalogue of fossil bees, 
the authors did not have the opportunity to re-examine the American species, so 
that the taxonomic aspects of these are not treated in as great a depth as are the 
European ones. In the main the authors have had to rely on the original authors* 
descriptions and figures for the American species except where the specimens were 
deposited in a European institution. 

Since this work was carried out some years ago, references to various institutions 
may be badly out-of-date, especially the war-damaged European museums. It has 
not, however, been the principle of the editor to undertake any significant alteration 
to the typescript as it stood. The taxonomic work as published is in two parts, 
the Apoidea excluding the Apidae (Section IV), and the Apidae (Section V). The 
division appears to have been made in this way because it most nearly represents 
the division between the New World and Old World fossil bees, and allows for 
different treatments of the two parts. 

The textfigures and plates had been made but have disappeared and were not 
amongst the effects of either of the authors. Fortunately the British Museum 
(Natural History) still retained negatives of many of the photographs and these 
have been reprinted for the plates. No attempt has been made to redraw the missing 
textfigures, since it is not known what they were or what they were intended to show. 
New photographs have been taken of specimens that are deposited in the British 
Museum (Natural History). 

Professor Zeuner was a Research Associate of the British Museum (Natural 
History), which position he had held since 1934 until his death. His obituary was 
published in 1963 in Nature, Lond. 200 : 1263. An obituary for Dr Manning 
appeared in 1966 in Proc. R. ent. Soc. Lond. (C) 31 : 62. 


The study of fossil bees, like that of any other group, requires a reasonably satis- 
factory system of classification. It may not always express fundamental relation- 
ships, but it must be consistent with the morphological evidence available. Michener 
(1944, 1965) attempted a revision of the families, subfamilies and tribes of bees of 
the world and of the North American genera in particular - a formidable task. From 
its very magnitude it was clear that the result would not completely satisfy the 
author himself, nor could it entirely agree with all the considered opinions of all bee 
taxonomists. His groups, however, well express relationships that have for long 
been obscure, and the inclusion of the bee families in the Sphecoidea has much to 
recommend itself. For these reasons, Michener's classification of the bees has been 
adopted in the present monograph. 

In the past, the lack of uniformity of the use of a terminology for their morphology 
has been a serious difficulty in the study of bees. Fortunately Snodgrass (1935) 
and more recently Michener (particularly 1944) have devoted considerable time and 
effort to clarifying this unhappy state of affairs, and we have taken advantage of 
the results of the labours of these authors. Moreover, the often excellent preser- 
vation of the fossil forms makes it necessary to treat them as if they were Recent 


Specimens, and to make detailed reference to morphological characters. This is 
particularly true, for instance, of specimens from Rott which show such intimate 
features as wax mirrors and parts of the alimentary canal. A modern terminology, 
therefore, is imperative. The terms used in this monograph for the structural 
features of the hind legs are defined later. 

With regard to wing venation the system of Ross (1936) has been adopted, together 
with the use of the following terms : submarginal cells, submarginal cross veins 
(i.e. 2nd abscissa Rs, ist r-m, 2nd r-m) and recurrent veins (ist m-cu and 2nd m-cu). 
The term 'intercubitus' has become obsolete with the rejection of the use of the 
term 'cubital cells' for the submarginal cells. The very convenient term 'basal 
vein', however, is retained for the free (i.e. not fused) sections of both M and Rs, 
which run from opposite directions towards each other before they unite to form a 
vein Rs + M. 

In addition, the term 'bomboid' is frequently used in the discussions that follow. 
This is not synonymous with ' Bombus-like' but is intended to suggest that the 
characters in question are merely reminiscent of those of the tribe Bombini. Bom- 
boid forms need not be closely related to Recent Bombini, though perhaps derived 
from either the ancestral form of the Bombini or, just as probably, from the ancestral 
form from which both the Bombini and the other Apidae arose. The last-mentioned 
possibility might imply that the Apini and Meliponini are not descended from the 
Bombini. The term 'apoid' is used in a somewhat similar sense in its appropriate 

Other terms, important because of their evolutionary significance, are those 
defining certain structures of the hind legs. The most essential are the following : 
(i) tibial comb, (2) tibial rake, or pollen rake, (3) tibial spur, (4) auricle, (5) depstum, 
and (6) basitarsal brush. 

The TIBIAL COMB^ is a structure much like a comb, situated on the outer side of 
the apex of the tibia, above the tibio-basitarsal joint. It consists of strong hairs, 
fused at the base, which usually project backwards and downwards. 

The TIBIAL RAKE is the dense row of spines or spine-like hairs along the apical 
edge of the inner side of the tibia. 

The TIBIAL SPURS are the two large and sharp spines which are jointed to the 
tibia. They protrude from the lower apical margin of the hind tibia. In the normal 
walking position, with the hind legs slightly turned outwards, they have to be looked 
for underneath in a seemingly forward position. If two spurs are present they are 
distinguished as the inner and outer hind tibial spurs. The inner one is often serrate. 
If only one is present, it may be either the outer or the inner, and one can be dis- 
tinguished from the other by its position relative to the apical rim of the tibia and 
by the presence or absence of serration, sometimes expanded into a comb-like 
structure. Thus a single spur with one edge serrate, occupying a relatively deep, 

1 The honey bee and the bumble bee lack this comb, but nevertheless have one or two hairs situated 
on the outer apical face of the corbicula. The derivation of these hairs is not known. Their position, 
however, precludes them from being direct derivatives of the tibial comb. They have not been noticed 
in fossil Apinae but might easily have been overlooked. They are used for stabilizing the load of 


inner position, is assumed to be the inner tibial spur, and a straight spur without 
serration, occupying a more marginal position, is assumed to be the outer. 

The AURICLE is the more or less broadened upper tip of the basitarsus which lies 
below the distal end of the tibia. It is like a projecting lip neatly fitting the com- 
plementary shape of the tibial apex, and in Recent Apis it is set with minute stud- 
like eminences. Often both the lip and tibial apex have been jointly termed the 
auricle, a practice not followed here. 

Depstum is a term used for an angularity of the basitarsus which is sometimes 
present where no true auricle is developed. This is a primitive condition, although 
some bees (e.g. Meliponini) do not even have a depstum in the modern species. 

Lastly, the term basitarsal brush is reserved for the neatly aligned rows of 
bristles present on the inner surface of the basitarsus of the Apini. It is not used for 
the haphazard arrangement of hairs found in other groups. 


For the study of the fossil Apidae a fair amount of material was available in the 
British Museum (Natural History), the total number of specimens being 52. The 
Meliponini, for instance, lacked only the specimens from the Sicilian amber which 
had been well described and illustrated by Tosi as long ago as 1896. The Apini 
lacked only representatives of the bees from Randecker Maar, mainly in the Arm- 
bruster Collection, but these were examined by one of the authors. It was only 
in the Bombini that the Museum collection really lacked material, due to the 
extreme paucity of the fossils of this tribe. Fortunately the most important 
representative, Bombus proavus Cockerell, has been well photographed and described 


Furthermore, during 1950 and again in 195 1, both authors travelled independently 
on the European continent studying fossil Apidae. Collections in Holland, Germany 
and Switzerland were examined. One of us had carried on this study of continental 
material for a number of years before it was decided to continue the work jointly. 

Whilst fossil Apoidea are plentiful in Europe, the absence of material belonging to 
families of bees other than the Apidae is disappointing. This material is chiefly in 
American museums and must await discussion by someone from there. Our 
attempts to obtain information or specimens from the relevant museums remained, 
unfortunately, without significant success. Below are given the names of the more 
important collections of fossil Apidae or museums where such material is to be 

Armbruster Collection. In the sediments of a small volcanic lake at Randeck, 
Wiirttemberg, honey bees have been found by numerous collectors. Possibly the 
oldest known specimens are those in the Oscar Fraas Collection, Stuttgart. In 
1926, however, William Scheuthle of Goeppingen began his search for fossil honey 
bees, and just over a year and a half later was assisted by Professor L. Armbruster. 
When the former died in November 1928, substantial discoveries had already been 
made and the collection eventually passed entirely into the possession of Professor 


Armbruster. Other collections believed to contain Randecker Maar honey bees 
are those of Eduard Scheer of Goeppingen, Bernard Hauff of Holzmaden, Karl 
Schempp of Brucken, and Pfarrer Hermann of Holzmaden. 

Bauckhorn Collection. See Siegburg an der Lahn (p. i6o). 

BuRSEY Collection. The collection of amber fossils formerly belonging to Mr 
Maurice Bursey of Surbiton, Surrey, now in the Museum National d'Histoire 
Naturelle, Paris, contains the important specimen E. {Electrapis) apoides Manning 
(p. 227). 

CocKERELL COLLECTION. The late Professor T. D. A. Cockerell of the University 
of Colorado collected and studied fossil bees. His main collections were made from 
the Miocene lake deposits of Florissant, Colorado, particularly during the expedition 
of 1906-7, when he was accompanied by his wife, Dr W. M. Wheeler and Mr S. A. 
Rohwer. His collections of fossil bees appear to have been few, and are now in 
the University of Colorado Museum, Boulder, Colorado, and the American Museum 
of Natural History. Although more of his other fossil insects are in the British 
Museum (Natural History), the only bee from his collection to be found there is 
Anthophora melfordi Cockerell. 

Danzig, Poland : Westpreussisches Provinzial-Museum. Many Baltic amber 
specimens were to be found here, but the whereabouts of the collection is not known 
at the present time. 

Karlsruhe i.B., Germany : Badische Landessammlung fiir Naturkunde, 
Erbprinzenstrasse 13. Contains some of the Oeningen and Radoboj material 
described by Oswald Heer. The building itself is in ruins, but the cellars are still 
packed with material and it is possible, therefore, that some further types, at present 
untraceable, will be found. 

Klebs Collection. From time to time Baltic amber specimens were formerly 
sold to private collectors and to museums, and it appears that this dispersal was 
made by the firm of Messrs Stantien & Becker, who traded the amber and who later 
were taken over by the Prussian State Amber Works. Messrs Stantien & Becker, 
in their day, placed the amber fossils in charge of Richard Klebs (1850- 191 1) who 
was, in later years, state geologist to the Prussian Geological Survey. A set of 346 
specimens, including the holotype of E. [Roussyana] proava Menge (p. 236), was 
acquired by the British Museum (Natural History) from Messrs Stantien & Becker 
in 1892, but since the specimens bear labels marked 'R. Klebs, Museum Stantien & 
Becker', this collection has usually gone under the name of 'Klebs Collection'. 
(See also p. 238.) 

KoENiGSBERG, East Prussia, U.S.S.R. : Geologisch-Palaontologisches Institut 
und Museum der Universitat. The most important types and described specimens 
of Baltic amber bees were included in this collection. They are now housed in the 
Museum of the Humboldt University, Berlin. 

Mainz (Rhein), Germany : Naturhistorisches Museum der Stadt Mainz. Contains 
a large collection of insect remains from the Hydrohia-Mm^sione of the district around 


London, British Museum (Natural History). See under Klebs Collection, 
Swinhoe, R. C. J., Luke Thomas Collection and Krantz, F. This collection includes 
some 52 specimens, including the types of Heyden (1862), and others from the 
Baltic amber, Rott am Siebengebirge, Florissant, Bottingen, East African copal, 
Burmese copal and South Australia. 

Marseille, France : Musee. According to Armbruster (1938) and Roussy (per- 
sonal communication), the specimen of 'Apis' aquitaniensis de Rilly (No. 5979) 
(p. 250) is in this collection, and Meunier (1915) states that a specimen of Antho- 
phorites mellona Heer (p. 196) is also in it. No confirmation, however, can be ob- 
tained after repeated enquiries. The Marseille Museum is rich in fossil insects 
from the Tertiary of southern France. 

RoussY Collection. This is the private collection of Monsieur Louis Roussy, 
Aigle, Switzerland, and contains mainly Baltic amber specimens, including Eledrapis 
[Roussy ana) palmnickenensis (Roussy), p. 233. 

ScHEELE Collection. This important collection of several thousand specimens 
of Baltic amber has been purchased by the Geologisches Staatsinstitut, Hamburg. 
Several important specimens of Eledrapis are included. 

SiEGBURG AN DER Lahn, Germany : Stadtisches Heimatmuseum. This museum 
contains the valuable collection of fossil insects made by Hugo Bauckhorn. It is 
particularly rich in specimens from Rott, previously studied by Meunier and 

Statz Collection. Fossil insects from Rott am Siebengebirge have been 
collected for over a century and are to be found in many museums and institutions. 
The collection made by the late Georg Statz of Cologne has become famous for the 
number and quality of its specimens. Moreover, Statz described and figured large 
numbers of his specimens. His collection includes several dozen fossil bees some of 
which have been well described and play an important role in the reconstruction of 
the phylogeny of the group. It is much to be regretted that this collection has been 
removed to Algiers, where it is in the hands of the collector's daughter. 

Stuttgart, Germany : Wiirttembergische Naturaliensammlung. This collection 
includes the material from Boettingen Swabian Alb, a locality situated not far from 
Randeck. One of us, while studying the fauna of this area, discovered a fossil 
swarm of honey bees, and casts were made of some of the bodies comprising it. 
Unfortunately it is believed that the original specimens were destroyed during the 
war, but the casts are still preserved in the British Museum (Natural History). 

Swinhoe Collection. Specimens of insect inclusions in the dark and pale 
varieties of Burmese amber from the Hukong Valley were collected by R. C. J. 
Swinhoe and presented to Professor T. D. A. Cockerell. Some were later presented 
by Cockerell to the British Museum (Natural History). Only the pale Burmese 
amber is known to contain bees. 

Timon-David Collection, Marseille. This collection consists of fossil insects 
from Camoins (Bassin de Marseille) and other French localities, and includes the 
fossil bee Halictus ruissatelensis Timon-David (p. 164). 


Luke Thomas Collection. A number of specimens of fossil bees are from East 
African copal. These, and other insects, have been presented to the British Museum 
(Natural History) at various times by individual donors. The Luke Thomas 
Collection contains twelve pieces of copal including fossil bees belonging to the 
Mehponini. They were presented to the British Museum (Natural History) in 
1945 by Col. H. Burrows. All are from the east coast of the mainland of Africa, 
facing Zanzibar. 

Washington, D.C, U.S.A. : United States National Museum ; Smithsonian 
Institution. Contains part of the collection, mainly from Florissant, described by 
Professor T. D. A. Cockerell. 

WiCKHAM Collection. The late Professor H. F. Wickham collected material 
from the Miocene shales of Florissant, Colorado, particularly at Wilson Ranch. 
This collection, which contains Andrena percontusa Cockerell (p. 167), is in Yale 

Zurich, Switzerland : Geologisches Institut und Museum der Eidgenossischen 
Technischen Hochschule. This collection includes much of the material, described 
by Oswald Heer, from Oeningen. 


The following list is arranged according to Michener (1944, 1965). His phylo- 
genetic groups and main diagnostic characters for famiUes, relevant subfamilies and 
tribes have been quoted, even though it is sometimes doubtful whether they are 
applicable to fossil forms. Indeed, only completely inapplicable characters have 
been omitted. 

In compiling the list a considerable number of specimens have been examined. 
In the case of unexamined material, no diagnoses of genera or species or systematic 
descriptions are given. These must await a study of the fossil material in American 
collections. Among such material, too, those specimens whose affinities have been 
stated in a definite way by earlier authors, but concerning which one might hold 
other views, have sometimes been accorded the benefit of the doubt, and noted 
under the generic name assigned by the most trustworthy author. The sections 
headed 'incertae sedis' therefore do not include specimens which are believed to be of 
such a nature that more information about them may substantiate their present 
suggested status. It is reserved for those which are too poorly preserved to be 
classifiable, and for vague records which cannot be substantiated by specimens. 

The publication containing the first valid name in accordance with the Inter- 
national Rules of Zoological Nomenclature has been marked with an asterisk (*). 

Superfamily APOIDEA Ashmead 1899 
Family GOLLETIDAE Bingham 1897 

Diagnosis. Labrum broader than long ; subantennal areas absent or at least 
reduced to small triangular spaces ; lower sides of clypeus not bent parallel to long 


axis of body ; subantennal sutures directed toward inner margins of antennal 
sockets ; facial foveae often present ; galeae short post-palpally and usually short 
pre-palpally, glossa short and often truncate or bifid, rarely round ; labial palpi 
short, segments similar but sometimes the first somewhat elongated and broadened ; 
pre-episternal sutures usually complete ; metanotum usually horizontal ; middle 
coxae, as seen from the outside, much shorter than the distance from their summits 
to posterior wing bases ; pygidial plate present or absent. 

Many of the genera are also distinguished from practically all other bees in 
having the posterior part of vein 2nd m-cu (2nd recurrent) of each forewing arcuate 
towards the apex of the wing ; the basal vein is not strongly curved. 

Remarks. No fossil CoUetidae have so far been found,^ though Cyrtapis, a 
genus of Halictidae, is according to Cockerell (1908a) in some respects reminiscent 
of the CoUetidae. 

Family HALICTIDAE Ashmead 1899 

Diagnosis. Labrum broader than long, but in some females its apex is produced 
to form a broad, acutely pointed process ; subantennal areas absent, though there 
is usually a single subantennal suture directed towards inner margins of antennal 
sockets ; lower sides of clypeus not bent back parallel to long axis of body ; facial 
foveae ordinarily absent ; galeae elongated pre-palpally, not post-palpally, the pre- 
palpal section tapering evenly to an acute base ; labial palpi with segments similar 
to each other, and usually cylindrical, rarely are any of the first three elongate and 
flattened. Glossa acute, usually short, without fiabellum ; pre-episternal sutures 
usually complete ; metanotum horizontal ; middle coxae with exposed parts much 
shorter than distance from summits to posterior wing bases ; basitibial plate usually 
present in females, except in parasitic genera ; pygidial plate present in females. 

Basal vein usually strongly curved ; jugal lobe of posterior wings much more than 
half as long as anal lobe. 

The most distinctive features are the long pre-palpal parts of the galeae, the 
usually strong arcuation or angulation of the ist abscissa of basal vein (vein M), and 
the normally complete pre-episternal suture. 

Distribution. Oligocene to Recent, 

Remarks. Apart from the genus Cyrtapis, which has coUetid affinities, only true 
Halictinae are known as fossils. By Oligocene times they appear to have been very 
modern in wing venation and probably in their general appearance also. 

Subfamily HALICTINAE Ashmead 1899 

Diagnosis. Supraclypeal area convex and protuberant if seen in profile ; clypeus 
itself longer than labrum and not protuberant much beyond level of supraclypeal 
area ; labrum in females commonly provided with a broad, pointed apical process ; 

2 But see Appendix, p. 255. 


antennal sockets not, or only a little, below middle of face ; pre-episternal suture 
conspicuous and complete ; scopa, when present, usually very extensive and found 
on posterior trochanters, femora, tibiae and basitarsi ; prepygidial fimbria usually 
divided by a longitudinal line without hairs or with only appressed hairs. 

First abscissa of basal vein (vein M) strongly arcuate and first submarginal cell 
ordinarily longer than third when, as is usually the case, three are present. 

Sometimes the glossa is elongated and variously modified ; the labial palpi are 
not elongated. 

Remarks. This large subfamily is only very moderately represented by fossils. 

Genus CYRTAPIS Cockerell 1908 

* 1908a Cyrtapis Cockerell : 339. 
1909b Cyrtapis Cockerell ; Cockerell : 80. 

Genotype. Cyrtapis anomalus Cockerell by monotypy. 

Remarks. According to Cockerell (1908a), 'the general appearance and the large 
stigma, marginal cell, and bent basal nervure are all as in the Halictines. The form 
of the second submarginal cell, with the first r.n. joining it before the middle, as well 
as the second r.n. with the double curve, suggest affinity with the CoUetines'. 
Cyrtapis may well have retained primitive elements, but because its evolution is 
towards the Halictines it is better to look upon it, for the time being at least, as a 
representative of the latter. Special mention of the type was made by Cockerell 

Cyrtapis anomalus Cockerell 1908 

♦1908a Cyrtapis anomalus Cockerell : 339. 
1909b C. anomalus Cockerell ; Cockerell : 80. 
1 93 1 Cyrtapis anomalus Cockerell ; Salt : 145. 

HoLOTYPE. Cockerell Collection, whereabouts unknown. 

Distribution. Oligocene : Florissant. 

Remarks. Mouth parts not known. In the circumstances, and because vein 
2nd m-cu (2nd recurrent) has a strong double bend, the species is considered to be a 
little anomalous. 

Genus HALICTUS Latreille 1804 

*i8o4 Halictus Latreille : 182. 
1935b Halictus Latreille ; Richards : 170. 
1938 Prohalictus Armbruster : 48, fig. 74 (type species : P. schemppi Armb.). 

Genotype. Apis quadricincta Fabricius, by subsequent designation of Richards 
(1935b : 170). 


Halictus ruissatelensis Timon-David 1944 

1943 Sphecodes Timon-David : 256. 
*i944 Halictus ruissatelensis Timon-David : 42. 

HoLOTYPE. Timon-David Collection, Marseille. 

Distribution. Oligocene : Bassin de Marseille. 

Remarks. Originally included by Timon-David (1943) in Sphecodes, but later 
(1944) placed, more correctly, in Halictus. We have not examined the specimen. 

Halictus florissantellus Cockerell 1906 

♦1906 Halictus florissantellus Cockerell : 43. 
1907 Halictus florisantellus Cockerell ; Handlirsch : 889. 

HoLOTYPE. Mus. Comp. Zool. Harvard, No. 2010 (= Scudder Coll. No. 921). 

Distribution. Oligocene : Florissant. 

Remarks. The name 'florisantellus' (Handlirsch 1907) is obviously a printer's 
error. Cockerell (1906) writes that only part of the wing venation is preserved, but 
'generic reference seems safe'. Approximately 6-5 mm long. 

Halictus miocenicus Cockerell 1909 

* 1 909a Halictus miocenicus Cockerell : 160. 
1931 Halictus miocenicus Cockerell ; Salt : 146. 

Holotype. Cockerell Collection, whereabouts unknown. 

Distribution. Oligocene : Florissant. 

Remarks. According to Cockerell, the generic position is probably correct. 

Halictus scudderiellus Cockerell 1906 

*i9o6 Halictus scudderiellus Cockerell : 43. 
1907 Halictus Scudderiellus Cockerell ; Handlirsch : 889. 

Holotype. Mus. Comp. Zool. Harvard, No. 2011 (= Scudder Coll. No. 1966). 

Distribution. Oligocene : Florissant. 

Remarks, Appears to be correctly placed. This bee is smaller than H. floris- 
santellus, being only 4-5 mm long. 

Halictus schemppi (Armbruster 1938) 

♦1938 Prohalictus schemppi Armbruster : 48, fig. 74. 
Holotype. Schempp Collection, Stuttgart- Weil. 
Paratypes. Three specimens, Armbruster Collection. 
Distribution. Miocene : Randeck. 


Remarks. Armbruster refers to one of his specimens as a true Halictus, stating 
(1938 : 48) that 'the nervulus is positioned quite normally'. There is no reason for 
believing the others are not conspecific. 

Family ANDRENIDAE Latreille 1802 

Diagnosis. Labrum broader than long, subantennal areas defined by two sub- 
antennal sutures beneath each antennal socket ; anterior lateral margins of clypeus 
usually concave ; lower sides of clypeus bent backwards more or less parallel to 
long axis of body (except in many species of Andrena) but clypeus not strongly 
protuberant ; facial foveae usually present ; galeae short pre-palpally and usually 
so post-palpally ; labial palpi short, the segments similar or first segment elongated 
and flattened, very rarely first two segments elongated ; glossa short or long, acute, 
without a flabellum ; pre-episternal sutures absent or present below scrobal sutures ; 
metanotum horizontal ; middle coxae short as seen externally, far shorter than 
distance from summits to posterior wing bases. Basitibial and pygidial plates 
present in females. 

Jugal lobe of posterior wing at least nearly three-quarters as long as anal lobe. 

The most characteristic feature of this family is the presence of defined sub- 
antennal areas. Unfortunately, in some species they lie below the bases of the 
antennae, covered with hairs, and even when uncovered only weak sutures may be 

Distribution. Eocene to Recent. 

Remarks. Only the subfamilies Andreninae and Panurginae are represented 
among the fossils. Andrena was well characterized as a genus by the Eocene. 

Subfamily ANDRENINAE Latreille 1802 

Diagnosis. Facial foveae large and usually covered with minute hairs giving a 
golden or reddish brown sheen in females, but in males foveae absent or virtually 
undefined and not clothed with minute hairs ; first flagellar segment rarely a little 
more than half as long as scape, usually much shorter ; maxillary palpi six- 
segmented ; segments of labial palpi cylindrical, similar to one another ; glossa 
usually short, acute ; pre-episternal sutures absent below scrobal sutures ; scopa 
present from coxae to basitarsi of posterior legs. 

Pterostigma moderate to large in size ; marginal cell longer than distance from 
apex to wing tip, and with the apex itself pointed on the costal margin of the wing 
or very narrowly rounded. 

Remarks. Most fossil members of this subfamily belong to the important genus 
Andrena, and two have been placed in the extinct genera Lithandrena said Pelandrena. 

Genus ANDRENA Fabricius 1775 

*i775 Andrena Fabricius : 376. 
1873 Biareolina Dours : 288 (type species : B. neglecta Dours). 
191 2 Andrena Fabricius ; Viereck : 613. 



Genotype. Apis helvola Linnaeus 1758, by subsequent designation of Viereck 
(1912 : 613). 
Distribution. Eocene to Recent. 

Andrena wrisleyi Salt 1931 
*i93i Andrena wrisleyi Salt : 141, fig. i. 

HoLOTYPE. Formerly in the Geol. Museum, Univ. Koenigsberg, but now in the 
Humboldt Museum, Berlin. 

Distribution. Eocene : Baltic amber. 
Remarks. Well figured and described. 

Andrena clavula Cockerell 1906 

*i9o6 Andrena (?) clavula Cockerell : 45. 
1907 Andrena (?) clavula Cockerell ; Handlirsch : 889. 

Holotype. Mus. Comp. Zool. Harvard, No. 2014 (= Scudder Coll. No. 6963). 

Distribution. Oligocene : Florissant. 

Remarks. Considered by Cockerell (1906) to be congeneric with A. sepulta. 
He remarked, however, that 'the shape of the abdomen is like that of a $ Ceratina, or 
possibly certain Halictines, but the venation does not agree with these'. 

Andrena grandipes Cockerell 1911 

*i9iia Andrena grandipes Cockerell : 73. 
193 1 Andrena grandipes Cockerell ; Salt : 145. 
1938 Andrena grandipes Cockerell ; Rodeck : 295. 

Holotype. Univ. Colorado Mus., No. 8599 (with counterpart). 

Distribution. Oligocene : Florissant. 

Remarks. Cockerell (1911a) states 'agrees closely with modern Andrena in 
most respects, but is peculiar for the very broad hind tibia and basitarsus, the small 
eyes and the long slender marginal cell'. 

Andrena hypolitha Cockerell 1908 

* 1908b Andrena hypolitha Cockerell : 328. 
1 93 1 Andrena hypolitha Cockerell ; Salt : 145. 
1938 Andrena hypolitha Cockerell ; Rodeck : 295. 

Holotype. Univ. Colorado Mus., No. 18736. 
Distribution. Oligocene : Florissant. 

Remarks. Differs from A. sepulta and A. clavula in various details of venation 
etc. It is not clear, however, how the abdomen differs from A. clavula. 


Andrena lagopus Latreille 1809 

*i8o9 Andrena lagopus Latreille : 151. 
1873 Biareolina neglecta Dours : 288. 
1913a Biareolina neglecta Dours ; Cockerell : 346. 

1930 Andrena lagopus Latreille ; Schmiedeknecht : 942. 

HOLOTYPE. Recent, whereabouts unknown. 

Fossil specimen. Possibly in the United States National Museum. 

Distribution. Oligocene : probably Florissant. Also Recent. 

Remarks. Schmiedeknecht, in identifying his specimen with a Recent species, 
mentions but one character, namely that the side of the stigma on the marginal cell 
'is conspicuously angulate or subdentate about the middle'. 

Biareolina Dours is a synonym of Andrena Fabricius, but since it possesses two 
submarginal cells, it is sometimes accorded separate generic status, a practice not 
followed by Michener (1944). Moreover, the reduction of the number of sub- 
marginal cells from three to two occurs frequently within the same species, for 
instance in Andrena, Halidus, Nontada and others. 

Andrena percontusa Cockerell 1914 

*I9I4 Andrena percontusa Cockerell : 640. 

1 93 1 Andrena percontusa Cockerell ; Salt : 145. 

HoLOTYPE. Wickham Collection, ? Peabody Museum, Yale. 

Distribution. Oligocene : Florissant. 

Remarks. Venation like A . sepulta and A . clavula, but a small species. Abdomen 
differs in shape from A. clavula. According to Cockerell (1914), the species can be 
distinguished from A . grandipes and A . hypolitha by the venation. 

Andrena sepulta Cockerell 1906 

*i9o6 Andrena sepulta Cockerell : 44. 
1907 Andrena sepulta Cockerell ; Handlirsch : 889. 

HoLOTYPE. Mus. Comp. Zool. Harvard, No. 2013 (= Scudder Coll. No. 14288). 
Distribution. Oligocene : Florissant. 

Remarks. Cockerell (1906) states 'the venation is not exactly like that of any 
modern species with which I have compared it but the differences are unimportant'. 

Andrena primaeva Cockerell 1909 

(PL I, fig. 2) 

1865 Drei Osmien ; Heer : 386 {partim). 
1876 Osmiae ; Heer : 43 {partim). 

1909c Three species of Osmia ; Cockerell : 316 [partim). 
* 1 909c Andrena (?) primaeva Cockerell : 316. 
1909c Osmia primaeva (Heer MS) ; Cockerell : 316. 
1 93 1 Andrena (?) primaeva Cockerell ; Salt : 145. 


HoLOTYPE, Eidgenossische Technische Hochschule, Zurich, No. SN.202, with 

Distribution. Miocene : Oeningen. 

Description and Remarks. Although Cockerell's references to Osmiae are, on 
the whole, difficult to interpret, there is no doubt whatever that the specimen he 
described as Osmia primaeva was the holotype. The insect is reasonably well 
preserved but the head is lacking. The chitin is dark brown in colour which may 
have been the colour of the original insect. The abdomen is plump and subglobose, 
but, since it is a millimetre broader than the thorax, the junction of the two gives the 
impression of a distinct constriction. 

The thorax has a large convex mesoscutum and apparently strong tegulae ; the 
other features are indistinct. 

The hind legs are fairly well preserved, the tibia and basitarsus being approxi- 
mately equal, their combined length being 3-5 mm ; the tibia is broad, apparently 
twice the width of the basitarsus, and both are copiously hairy ; the femur is as 
broad as the tibia. The tibia and basitarsus of the middle leg are not preserved in a 
natural position, and we are inclined to consider Cockerell's basitarsus as the tibia 
and his tibia as the femur. Both femur and tibia are of the same width, but the 
tibia is slightly longer. 

The wings are not preserved in their entirety owing to a crack in the rock. For- 
tunately, what is preserved is very distinct and includes the apical region of the fore 
and hind wings. 

Forewing : long, well-developed pterostigma ; marginal cell narrowing apically, 
and arising broadly from the pterostigma ; three submarginal cells, the first not 
completely preserved, the second rhomboid, receiving the first recurrent vein slightly 
distad of the middle and having its proximal cross vein slightly hooked at the base, 
the third submarginal cell at least twice as large as the second, only half as broad on 
the marginal cell as its own base, and receiving the second recurrent vein in its last 

Hindwing : radial sector and media continued to wing margin and cross-vein r-m 
only very slightly oblique. 

Cockerell's (1909) statement that 'all things considered the reference to Andrena 
seems reasonably assured' appears a fair conclusion. The venation of the hind wing 
and the rather plump body precludes the specimen from being referred to Halictus. 

Measurements. Length of thorax and abdomen (head lacking) 7 mm ; length 
of thorax 2-25 mm, width 3 mm ; length of abdomen 4-75 mm, width 4 mm. 

Andrena sp. indet. (Brischke) 

1886 Andrena Brischke : 278. 

1891 Andrena 'Bxischk.e ; Scudder : 686 (5915). 

1907 Andrena Brischke ; Handlirsch : 890. 

193 1 Andrena sp. Brischke ; Salt : 144. 

Distribution. Eocene : Baltic amber. 


Remarks. Among Menge's and Helm's collections, investigated by Brischke, 
were apparently two specimens generically determined as Andrena, but not described. 

Andrena sp. indet. (Motschulsky) 

1856 Andrena Motschulsky : 28. 

1 93 1 Andrena sp. Motschulsky ; Salt : 144. 

Distribution. Eocene : Baltic amber. 

Remarks. Motschulsky merely records the occurrence of the genus. 

Genus LITH ANDRENA Cockerell 1906 

*i9o6 Lithandrena Cockerell : 44. 
1909b Lithandrena Cockerell ; Cockerell : 81. 

Genotype. Lithandrena saxorum Cockerell 1906 by monotypy. 

Remarks. According to Cockerell, 'a genus of Andrenidae, allied to Andrena. 
It differs from Andrena and Nomia in the second r.n. which is strongly bent in its 
upper part, straight but oblique below ; from Andrena alone it differs in the propor- 
tions of the submarginal cells ; and from Nomia also by the tip of the marginal cell, 
which is pointed, and a little away from the costae. The general appearance is that 
of an Andrena, but it cannot be referred to this or any other genus . . .'. Special 
mention of the type was made by Cockerell (1909b). 

Lithandrena saxorum Cockerell 1906 

*i9o6 Lithandrena saxorum Cockerell : 44. 
1907 Lithandrena saxorum Cockerell ; Handlirsch : 889. 
1909b Lithandrena antiquorum Cockerell ; Cockerell : 80. 

HoLOTYPE. Mus. Comp. Zool. Harvard, No. 2012 (= Scudder Coll. No. 8219). 

Distribution. Oligocene : Florissant, 

Remarks. Length 8-6 mm. Venation of forewing fully described (Cockerell 
1906). Head as wide as thorax, black ; flagellum stout ; abdomen light in colour 
with a broad entire dark band on each segment, legs hairy. 

Genus PELANDRENA Cockerell 1909 

*i909a Pelandrena Cockerell : 159. 
1909b Pelandrena Cockerell ; Cockerell : 81. 

Genotype. Pelandrena reducta Cockerell 1909a by monotypy. 

Remarks. Allied to Andrena but with two submarginal cells on the anterior wing. 
Special mention of the type was made by Cockerell (1909b). The reason for the 
erection of this genus is not clear. It may be based only on the two submarginal 
cells, in which case its generic status is weak. 


Pelandrena reducta Cockerell 1909 

* 1 909a Pelandrena reducta Cockerell : 160. 
1909b Pelandrena reducta Cockerell ; Cockerell : 81. 
1931 Pelandrena reducta Cockerell ; Salt : 146. 
1938 Pelandrena reducta Cockerell ; Rodeck : 298. 

HoLOTYPE. Univ. Colorado Mus., No. 18741. 
Distribution. Oligocene : Florissant. 

SubfamUy PANURGINAE Leach 1815 

Diagnosis. Facial foveae, when present, much smaller than in Andreninae, not 
clothed with fine hairs, and frequently distinct in males as well as in females ; labial 
palpi frequently with first segment much elongated ; maxillary palpi usually six- 
segmented (reduced in some species of Perdita). Scopa primarily confined to pos- 
terior tibiae and basitarsi. 

Pterostigma usually rather large ; marginal cell, which is normally not longer 
than distance from its own apex to apex of wing, mostly obliquely truncate apically 
but sometimes merely bent away from wing margin. 

Remarks. Only one fossil genus belongs to this subfamily. 

Genus LIBELLULAPIS Cockerell 1906 

*i9o6 Libellulapis Cockerell : 42. 
1909b Libellulapis Cockerell ; Cockerell : 80. 

Genotype. Libellulapis antiquorum Cockerell 1906 by monotypy. 

Remarks. Cockerell (1913a), when discussing Libellulapis wilmattae, stated, 'A 
reconsideration of the genus leads me to place it in the Panurgidae where, by reason 
of certain features of the venation, it appears to stand rather near to Panurgus'. 
Eyes curiously prominent. Special mention of the type was made by Cockerell 

Libellulapis antiquorum Cockerell 1906 

*i9o6 Libellulapis antiquorum Cockerell : 42. 

1907 Libellulapis antiquorum Cockerell ; Handlirsch : 889. 

1909a Libellulapis antiquorum Cockerell ; Cockerell : 80. 

1938 Libellulapis antiquorum Cockerell ; Rodeck : 297. 

HoLOTYPE. Mus. Comp. Zool. Harvard, No. 2009 (= Scudder Coll. No. 9061). 

Paratype. Univ. Colorado Mus., No. 18740 (= Scudder Coll. No. 8560). 

Distribution. Oligocene : Florissant. 

Remarks. Cockerell (1906) considered this species different from Halictoides 
maurus (Cresson), Hesperapis rhodoceratus (Cockerell) and Parandrena andrenoides 
Cresson in points of venation, and believed that it did not agree with Diandrena or 



Libellulapis wilmattae Cockerell 1913 

♦1913a Libellulapis wilmattae Cockerell : 344, figs 2, 3. 
1931 Libellulapis wilmattae Cockerell ; Salt : 146. 
1938 Libellulapis wilmattae Cockerell ; Rodeck : 297. 

HoLOTYPE. U.S. Nat. Mus., No. 58688. Counterpart, Univ. Colorado Mus., 
No. 18742. 
Distribution. Oligocene : Florissant. 

Remarks. Cockerell (1913a) states, 'Broad head and prominent eyes as in some 
Panurgi, pygidial plate as in Dasypoda' . 

Family ANDRENIDAE incertae sedis 
Andrenidae gen. et sp. indet. (Handlirsch) 

1907 (? Andrenidae) ; Handlirsch : 890. 

193 1 Andrenidae ? sp. (Handlirsch) ; Salt : 144. 

Material. Hofmuseum, Vienna. 

Distribution. Eocene : Baltic amber. 

Remarks. Hind leg with pollen. Head formation suggestive of Halidus or 

Andrenidae gen. et sp. indet. (Scudder) 

1 88 1 Andrenidae ; Scudder : 290. 

1883 Andrenidae ; Scudder : 280. 

1891 (Andrenidae) (several) ; Scudder : 682 (5872). 

1907 (Andrenidae) (mehrere) Scudder ; Handlirsch : 890. 

Distribution. Oligocene : Florissant. 

Remarks. Specimens appear to be poorly preserved. 

Family MELITTIDAE Dumeril 1821 

Diagnosis. Labrum broader than long ; subantennal areas absent ; subantennal 
sutures directed, except sometimes at their upper ends, towards inner margins of 
antennal sockets ; lower sides of clypeus not, or only a little, bent backwards ; 
clypeus not protuberant. Facial foveae absent ; galeae short pre-palpally and 
usually so post-palpally ; labial palpi with segments similar to one another, cylin- 
drical ; glossa short or long, acute ; pre-episternal and scrobal sutures usually 
absent, the former always absent below the latter ; metanotum horizontal or sub- 
horizontal ; middle coxae short as seen externally, much shorter than distance 
from their summits to posterior wing bases ; scopa confined to posterior tibiae and 
basitarsi, although trochanters and femora are hairy ; basitibial plates and pygidial 
plate present in female. 

Distribution. Eocene to Recent. 


Remarks. The subfamilies Ctenoplectrinae and Melittinae only are represented 
among the fossils. 

Subfamily CTENOPLECTRINAE Cockerell 1920 

Diagnosis. Scrobal and upper parts of pre-episternal sutures are present, 
though of doubtful value for fossil forms ; inner hind tibial spur much broadened 
basally and throughout its length provided with a comb of very many long fine 
teeth ; posterior basitarsi of female nearly as broad as tibiae ; pygidial plate of 
male absent, that of female narrow apically, greatly broadened basally. 

Pterostigma hardly twice as long as broad ; marginal cell considerably longer than 
distance from its apex to wing tip and bent gradually away from wing margin for 
nearly half length of cell ; two submarginal cells ; jugal lobe of posterior wing about 
half as long as anal lobe. 

Remarks. Among Recent forms this subfamily contains only the genus Cteno- 
pledra found from Australia and Africa to China. This genus, however, is not yet 
known in the fossil state. On the other hand, the subfamily is represented by two 
fossil genera Ctenoplectrella and Glyptapis. Moreover, Cockerell (igogd) maintained 
that the genera included in this subfamily are 'so remote from the modern members 
of that group that they at least form a distinct subfamily, Glyptapinae'. That 
group', however, refers to the Megachilidae to which Ctenoplectra, and hence 
Ctenoplectrella etc., was considered to belong. Therefore until the affinities of the 
Ctenoplectra group, both fossil and Recent, have been more clearly defined, it is not 
considered advisable to create a new subfamily. 

Genus CTENOPLECTRELLA Cockerell 1909 

♦1909c Ctenoplectrella Cockerell : 314. 
igogd Ctenoplectrella Cockerell ; Cockerell : 19. 

Genotype. Ctenoplectrella viridiceps Cockerell 1909 by monotypy. 

Remarks. This genus appears to be closely related to Glyptapis. Both genera 
have Ctenoplectra as 'their nearest relative in the modern fauna'. C. viridiceps was 
first mentioned by Cockerell (1909c) with some characters, but its full description 
followed in a later paper (igogd). [See also Appendix, p. 255.] 

Ctenoplectrella dent at a Salt 1931 

♦1931 Ctenoplectrella dentata Salt : 139. 

HoLOTYPE. Formerly in the Geological Museum, University of Koenigsberg, now 
in the Humboldt Museum, Berlin. 

Distribution. Eocene : Baltic amber. 

Remarks. Salt considers it possible that this specimen is the male of Cteno- 
plectrella viridiceps Cockerell and gives a good description and figures. 


Ctenoplectrella viridiceps Cockerell 1909 

♦1909c Ctenoplectrella viridiceps Cockerell : 314. 
i909d Ctenoplectrella viridiceps Cockerell ; Cockerell : ig, fig. 14. 
1 93 1 Ctenoplectrella viridiceps Cockerell ; Salt : 145. 

HoLOTYPE. Formerly in the Geological Museum, University of Koenigsberg, now 
in the Humboldt Museum, Berlin. 

Distribution. Eocene : Baltic amber. 

Remarks. Female, see above. This species was a small, black, stout bee, like 
Glyptapis, hardly 5 mm long. 'The eyes were bare and the sting visible'. Full 
description given by Cockerell (igogd). 

Genus GLYPTAPIS Cockerell 1909 

* 1 909c Glyptapis Cockerell : 314. 
igogd Glyptapis Cockerell ; Cockerell 113. 

Genotype. Glyptapis mirahilis Cockerell, by original designation. 

Remarks. This genus, together with Ctenoplectrella, is referred by Cockerell 'to 
a group from which the Megachiloids sprang'. The genus was mentioned (1909c) 
in connection with G. mirahilis, but it was not described until later (i909d). 

Glyptapis fuscula Cockerell 1909 

*i909d Glyptapis fuscula Cockerell : 17, fig. 12. 
193 1 Glyptapis fuscula Cockerell ; Salt : 145. 

HoLOTYPE. Formerly in the Geological Museum, University of Koenigsberg, now 
in the Humboldt Museum, Berlin. 

Distribution. Eocene : Baltic amber. 

Remarks. Cockerell (i909d) mentions that the ventral side of the abdomen is 
clearly seen to be furnished with strong bristles, the beginning of a ventral scopa. 
As distinctive characters of the species he gives the very dark wings and the large 

Glyptapis mirabilis Cockerell 1909 

* 1 909c Glyptapis mirabilis Cockerell : 314. 
igogd Glyptapis mirabilis Cockerell ; Cockerell : 14, figs 9, 10. 
193 1 Glyptapis mirabilis Cockerell ; Salt : 146. 

HoLOTYPE. Formerly in the Geological Museum, University of Koenigsberg, now 
in the Humboldt Museum, Berlin. 

Distribution. Eocene : Baltic amber. 

Remarks. The genotype, selected by Cockerell (i909d), is actually the monotype, 
since the genus and species were first mentioned (1909c) together, with a few 


descriptive words. The specific characters are the dark brown stigma and veins, 
the length of the marginal cell (over i -2 mm) , the entirely black body and the punctate 

Glyptapis neglect a Salt 1931 
*i93i Glyptapis neglecta Salt : 136, fig. i. 

HoLOTYPE. Formerly in the Geological Museum, University of Koenigsberg, now 
in the Humboldt Museum, Berlin. 

Distribution. Eocene : Baltic amber. 

Remarks. Well described and figured by Salt. This is the largest known 
Glyptapis from the Baltic amber. 

Glyptapis reducta Cockerell 1909 

♦igogd Glyptapis reducta Cockerell : 18, fig. 13. 
1 93 1 Glyptapis redticta Cockerell ; Salt : 146. 

HoLOTYPE. Formerly in the Geological Museum, University of Koenigsberg, now 
in the Humboldt Museum, Berlin. 

Distribution. Eocene : Baltic amber. 

Remarks. Wings hyaline. Cockerell (i909d) regards the ferruginous stigma 
and veins and the length of the marginal cell (0-935 mm) as diagnostic. 

Glyptapis reticulata Cockerell 1909 

*igo9d Glyptapis reticulata Cockerell : 16, fig. 11. 
193 1 Glyptapis reticulata Cockerell ; Salt : 146. 
1969 Glyptapis reticula Cockerell ; Kellner-Pillault : 521. 

HoLOTYPE. Formerly in the Geological Museum, University of Koenigsberg, now 
in the Humboldt Museum, Berlin. 

Distribution. Eocene : Baltic amber. 

Remarks. Differs from G. mirahilis chiefly in the metallic crimson tints on the 
abdomen and the reticulate mesothorax. 

Subfamily MELITTINAE Dumeril 1821 

Diagnosis. Inner hind tibial spur unmodified ; posterior basitarsi more slender 
than tibiae (except in male Haplomelitta) and shorter than tibiae ; distitarsi unusually 

Pterostigma slender, more than twice as long as broad, its margins basad of vein 
are very rarely converging basally ; marginal cell longer than distance from its own 
apex to wing tip, the apex itself very narrowly rounded on wing margin and bent 
away from the latter, if at all, only apically ; three submarginal cells, second much 


the smallest, the first being the largest ; jugal lobe of posterior wing nearly half as 
long as anal lobe (rarely three-quarters). 

The species of this subfamily are of moderate size, and black. The abdomen is 
provided with pale pubescent fasciae, giving the appearance of certain species of 
Halictus and Andrena. 

Remarks. This subfamily is represented, among fossils, only by the genus 


Genus MELITTA Kirby 1802 

*i8o2 Melitta Kirby : 130-134. 
1935b Melitta Kirby ; Richards : 172. 

Genotype. Melitta tricincta Kirby 1802, selected by Richards (1935 : 172). 

Melitta willardi Cockerell 1909 

♦19096 Melitta willardi Cockerell : 393. 
193 1 Melitta willardi Cockerell ; Salt : 146. 
1938 Melitta willardi Cockerell ; Rodeck : 297. 

HoLOTYPE. Univ. Colorado Mus., No. 18737. 

Distribution. Oligocene : Florissant. 

Remarks. Cockerell (i909e) writes, 'Venation normal for Melitta, except that 
the upper segment of the basal nervure is shorter' ; also 'the reference of this insect 
to Melitta seems safe'. 

Family MELITTIDAE incertae sedis 

Dasypoda (s.l.) sp. indet. (Menge) 

1856 Dasypoda Menge : 26. 

1891 Dasypoda (vie.) Menge ; Scudder : 696 (6005). 

1907 Dasypoda (vie.) Menge ; Handlirsch : 890. 

1 93 1 Dasypoda vie. Menge ; Salt : 144. 

Material. Whereabouts unknown. 

Distribution. Eocene : Baltic amber. 

Remarks. An 'Andrenida, Grabwespen' was reported (Menge 1856) to resemble 
Dasypoda but 'was distinguished from it by the hind basitarsus having a blunt 
lateral process in front of the joint with the second tarsal segment, and the hind leg 
being longer than the abdomen' (transL). The mandibles had a broad tip with 
three teeth and the lancet-shaped tongue had labial palps protruding. Two speci- 
mens existed, one clearly visible, the other covered with 'schimmel'. 

Family MEGACHILIDAE Latreille 1802 

Diagnosis. Labrum longer than broad, widened basally to form a long line of 
articulation with clypeus ; subantennal areas absent ; subantennal sutures directed 


towards outer margins of antennal sockets ; lower sides of clypeus not bent back- 
wards parallel to long axis of body ; facial foveae absent ; first flagellar segment 
much shorter than scape ; galeae short pre-palpally, long post-palpally ; labial palpi 
with first two segments elongated and flattened, sheath-like, first segment usually 
shorter than second ; glossa linear, possessing a flabellum, pre-episternal and 
scrobal sutures considerably reduced or absent ; metanotum usually vertical, 
sometimes horizontal ; middle coxae elongate, at least half, and usually more than 
half, as long as distance from summits to hindwing bases ; basitibial plates absent ; 
scopa, when present, confined to abdominal sterna ; pygidial plate absent, except 
in Lithurge. 

Pterostigma small ; submarginal cells two, the second as long, or nearly as long, 
as the first ; marginal cell pointed or narrowly rounded at apex, which is usually a 
little bent away from costal margin of wing and nearer to wing tip than to base of 

The most distinguishing characters are the position of the scopa, the form of the 
labrum and the direction taken by the subantennal sutures. 

Distribution. Eocene to Recent. 

Remarks. The subfamilies Lithurginae and Megachilinae are known as fossils, 
the latter being well represented in the Oligocene. 

Subfamily LITHURGINAE Newman 1834 

Diagnosis. Face of female elevated below antennal sockets ; epistomal suture 
in most part absent ; mandibles in both sexes short and robust, tridentate, the 
middle tooth longer and more elevated than the others ; metanotum about one- 
eighth as long as scutellum ; coarse spiculae of tibiae arranged in two longitudinal 
rows on fore and middle legs, scattered on posterior legs ; posterior basitarsi slender, 
parallel-sided, about as long as posterior tibiae ; second abdominal tergum unusually 
small, posterior margin broadly rounded ; terga two to seven of male exposed and 
unmodified ; male with pygidial plate, female with slender remnant thereof in the 
form of large spine, flattened dorsally. 

Jugal lobe of posterior wing about three-quarters as long as anal lobe. 

Remarks. Only one genus, Lithurge, is known to which both Recent and fossil 
forms belong. 

Genus LITHURGE Latreille 1825 

♦1825 Lithurge Latreille : 463. 
1827 Lithurgus Berthold : 467. 

Genotype. Centris cornuta Fabricius 1787 by monotypy. 

Lithurge adamitica (Heer 1865) 

^1865 Apis adamitica Heer : 386, fig. 287. 
1867 Apis adamitica Heer ; Heer : 4, pi. 3, fig. 11 


1872 Apis adamitica Heer ; Heer : 473, fig. 287. 

1876 Apis adamitica Heer ; Heer : 43, fig. 287. 

1879 Apis adamitica Heer ; Heer : 412, fig. 329. 

1891 Apis adamitica Heer ; Scudder : 688 (5936). 

1907 Apis adamitica Heer ; Handlirsch : 892. 

1909c Lithurgus adamiticus (Heer) ; Cockerell : 313. 

1915 Apis adamitica Heer ; v. Buttel-Reepen : 9, fig. i. 

1928 Apis adamitica Heer ; Wheeler : 97. 

1928 Lithurgus Berthold ; Wheeler : 97. 

193 1 Apis adamitica Heer ; Statz : 45. 

1938 'Apis adamitica' (Heer) ; Armbruster : 86. 

HoLOTYPE. Eidgenossische Technische Hochschule, Zurich, V.S.4. 

Other specimens, (i) Eidgenossische Technische Hochschule, Zurich, SN.200 ; 
(2) Badische Landessammlung fiir Naturkunde, Karlsruhe, Inv.1216 (Oe.95). 

Distribution. Miocene : Oeningen. 

Description and Remarks. Cockerell (1909c) considered this species to be 
somewhat more primitive than recent Lithurge, possibly a near subgenus. According 
to him it is separated from Apis by its possession of only two submarginal cells. 
Unfortunately the holotype, which we have studied, is not well preserved and we 
have not succeeded in recognizing the submarginal cells. Armbruster (1938) who 
contributed some original remarks on it did not consider it to belong to Apis. 

The holotype, indeed, is a somewhat elongated insect with a slender and pointed 
abdomen. The outlines of the wings are preserved but the venation is mostly too 
indistinct for satisfactory recognition. It is clear, however, that the species does 
not belong to the genus Apis, and also that the preservation of the specimen appears 
to have deteriorated since it was examined by Cockerell. 

The following points are presented as still being of interest. The head and thorax 
have preserved no recognizable diagnostic characters ; the colour of the body has 
darkened from Cockerell's 'warm, red-brown' ; the abdomen, as Cockerell stated, is 
'truncate basally, pointed apically' and, since basal intersegmental membranes are 
evident, may be somewhat distended ; legs are lacking ; wings are small in compari- 
son with size of insect, and wing venation shows only the following points. Strong 
costal vein ; straight and short basal vein, the anterior section of which is longest ; 
short, stubby, pointed marginal cell ; poorly developed pterostigma. The apex of 
the wing, marginal cells and recurrent nerves are no longer clearly evident ; the 
cross vein cu-a, however, can be detected slightly distal of the juncture of basal 
vein on cu. While these characters obviously do not give sufficient grounds for 
placing the specimen generically, we must recognize that Cockerell saw it while in a 
better state of preservation and accept his decision in this matter. 

A second specimen (Zurich, SN.200) has been studied and tentatively assigned to 
this species. Its head, thorax and abdomen (except the tip) are quite well preserved, 
but the wings are present only in traces and the legs are missing. A strong costal 
vein is present, and a marginal cell similar to Lithurge. 

A third specimen (Karlsruhe, Inv.1216 (Oe.95)) may also belong here. Abdomen 
and thorax alone are preserved. The former, however, is well segmented and 
pointed, and is very reminiscent of the holotype, from which it differs only slightly 


in size, i.e. length of abdomen lo mm, as compared with 8-5 mm for the holotype. 
The width of the abdomen is 4-25 mm and the overall length, as preserved, i.e. 
without head, is 13 mm. 

Finally, it is interesting to note that Heer's figure (1865 : fig. 287) differs in wing 
venation from his pi. 3, fig. 11 (1867), although both are presumably meant to repre- 
sent the same specimen. 

Subfamily MEGACHILINAE Latreille 1802 

Diagnosis. Face of female very rarely elevated below antennal sockets ; epi- 
stomal suture complete ; mandibles variousl}" toothed, apical tooth longest, preapical 
tooth not elevated ; tibiae not spiculate ; posterior basitarsi much shorter than 
tibiae ; second abdominal tergum large, with the posterior margin usually straight, 
so that base of abdomen lacks the compressed aspect of Lithurge ; abdomen of male 
with fewer unmodified sterna than in Lithurge ; pygidial area entirely absent. 

Jugal lobe of posterior wings less than half as long as anal lobe. 

Remarks. Two tribes, Anthidiini and Megachilini, are distinguished within this 
subfamily, both known as fossils. 

Tribe ANTHIDIINI Michener 1944 

Diagnosis. Posterior margin of scutellum usually vertical or even overhanging, 
at a distinct angle to the dorsal surface ; integumental maculation, at least in part, 
pale ; claws of female cleft or at least with an inner tooth. 

Inner margin of pterostigma from its base to the base of medial vein little, if at 
all, longer than width of pterostigma. 

Remarks. Two Recent genera are represented by fossils, and there is one extinct 
genus. [See also Appendix, p. 256.] 

Genus ANTHIDIUM Fabricius 1805 

*i8o5 Anthidium Fabricius : 364. 
1810 Anthidium Fabricius ; Latreille : 439. 

Genotype. Apis manicata Linnaeus 1758, selected by Latreille (1810 : 439). 

Anthidium mortuum (Meunier 1920) 

♦1920 Eucera mortua Meunier : 731, pi. i, fig. 5, text-fig. 4. 
1 93 1 Eucera mortua Meunier ; Salt : 145. 
1936 Anthidium mortuum (Meunier) ; Statz : 260, 285, 293, 294, pi. 13, fig. 38. 

Holotype. Heimatmuseum Siegberg (Lahn), Bauckhorn Collection. 

Distribution. Oligocene : Rott am Siebengebirge. 

Remarks. This specimen was first described as Eucera by Meunier (1920). 
Statz (1936), on re-examining the specimen, placed it in Anthidium. He insisted 
that the venation of the fossil form was identical with that of Anthidium, except for 


the 'nervulus' which is 'interstitial' in the fossil and 'postfurcal' in the Recent species. 
Since Statz was the more reliable worker of the two, we are placing this form in 
Anthidium. The possibility of a Eucera occurring in Rott should not, however, be 
wholly disregarded, since Statz is anxious to reinforce his identification by the 
ecological argument that Anthidium builds its nest in stalks and twigs, which agrees 
better with the Rott biotope than the steppe habitat of Eucera. Statz' (1936 : 260), 
however, states that Recent species of Eucera have the second recurrent vein meeting 
the second submarginal cell, whereas in Meunier's Eucera mortua it ends distad to 
the second submarginal cross vein as in the genus Anthidium. 

It is also necessary to point out that Statz' discussion of this form is misleading. 
He describes it (1936 : 285) under the heading of 'new discoveries', whilst his figure 
proves it to be Meunier's holotype. Three further disconnected paragraphs appear 
(1936 : 260, 293, 294) in which no mention is made of the fact that he had no new 
specimen. The identity with Meunier's holotype is proved not only by the general 
shape of the fossil, but also by the frayed margin of the right forewing. 

Anthidium exhumatum Cockerell 1906 

♦1906 Anthidium exhumatum Cockerell ■.38. 
1907 Anthidium exhumatum Cockerell ; Handlirsch : 888. 

Holotype. Mus. Comp. Zool. Harvard, No. 2003 (= Scudder Coll. No. 13709, 
with counterpart. No. 11388). 

Paratype. Scudder Coll. No. 8444. 

Distribution. Oligocene : Florissant. 

Remarks. According to Cockerell, 'not so typical an Anthidium as A. scudderi'. 
The paratype agrees with the holotype in the width of the marginal cell and the 
shape of the second discoidal. It differs in the strongly banded abdomen. He 
concludes, 'It appears to be a female and I have little doubt that it represents that 
sex in A. exhumatum'. 

Anthidium scudderi Cockerell 1906 

*igo6 Anthidium scudderi Cockerell : 38. 
1907 Anthidium scudderi Cockerell ; Handlirsch : 888. 

Holotype. Mus. Comp. Zool. Harvard, No. 2002 (= Scudder Coll. No. 11381). 
Distribution. Oligocene : Florissant. 

Remarks. Apparently a typical Anthidium which can be clearly separated from 
Megachile on points of wing venation. 

Genus DIANTHIDIUM Cockerell 1900 

*i9oo Dianthidium Cockerell : 412. 

Genotype. Anthidium curvatum Cockerell {nee Smith) - Dianthidium sayi 
Cockerell, by original designation (1900). 


Dianthidium tertiarium Cockerell 1906 

*i9o6 Dianthidium tertiarium Cockerell : 39. 
1907 Dianthidium tertiarium Cockerell ; Handlirsch : 888. 

HoLOTYPE. Mus. Comp. Zool. Harvard, No. 2004 (= Scudder Coll. No. 806). 
Distribution. Oligocene : Florissant. 

Remarks. This average-sized (8 mm) dark brown bee can be assumed to be 
correctly placed by Cockerell, in view of his intimate knowledge of the genus. 

Genus LITHANTHIDIUM Cockerell 1911 
♦igiib Lithanthidium Cockerell : 225. 

Genotype. Lithanthidium pertriste Cockerell 1911 by monotypy. 

Lithanthidium pertriste Cockerell 1911 

*i9iib Lithanthidium pertriste Cockerell : 225. 
1 93 1 Lithanthidium pertriste Cockerell ; Salt : 146. 
1938 Lithanthidium pertriste Cockerell ; Rodeck : 297. 

Holotype. Univ. Colorado Mus., No. 18738. 
Distribution. Oligocene : Florissant. 
Remarks. Only a single specimen known. 

Tribe MEGAGHILINI Latreille 1802 

Diagnosis. Claws of female simple ; no obtrusive pale integumental maculation ; 
abdominal pattern often present, resulting from bands of pale hairs ; posterior part 
of scutellum, except in some parasitic forms, not separated by a sharp angle from 
dorsal surface. 

Inner margin of pterostigma from its own base to base of radial vein much longer 
than width of pterostigma. 

Remarks. The Recent genera Heriades, Megachile and Osmia are represented by 

Genus HERIADES Spinola 1808 

*i8o8 Heriades Spinola : 7. 
1 8 10 Heriades Spinola ; Latreille : 439. 

Genotype. Apis truncorum Linnaeus 1758, selected by Latreille (1810 : 439). 
Remarks. According to Spinola's usage, the name Heriades is feminine. The 
masculine ending, however, is used here in conformity with modern usage. 


Heriades bowditchi Cockerell 1906 

*i9o6 Heriades bowditchi Cockerell : 41. 
1907 Heriades bowditchi Cockerell ; Handlirsch : 889. 
1938 Heriades bowditchi Cockerell ; Rodeck : 296. 

HoLOTYPE. Mus. Comp. Zool. Harvard, No. 2007 (= Scudder Coll. No. 13761). 
Paratype. Univ. Colorado Mus., No. 18739 (= Scudder Coll. No. 13436). 
Distribution. Oligocene : Florissant. 

Remarks. Differs from H. laminarum in the pronouncedly larger head and the 
banded abdomen. 

Heriades halictinus Cockerell 1906 

*i9o6 Heriades halictinus Cockerell : 40. 
1907 Heriades halictinus Cockerell ; Handlirsch : 889. 

HoLOTYPE, Mus. Comp. Zool. Harvard, No. 2006 (= Scudder Coll. No. 10564). 

Distribution. Oligocene : Florissant. 

Remarks. Resembles a small Halictus in superficial aspect, but its true affinity 
is indicated by wing venation. Differs from H. laminarum in its darker abdomen 
and smaller size. 

Heriades laminarum Cockerell 1906 

*i9o6 Heriades laminarum Cockerell : 40. 
1907 Heriades laminarum Cockerell ; Handlirsch : 888. 

HoLOTYPE. Mus. Comp. Zool. Harvard, No. 2005 (= Scudder Coll. No. 3062). 
Distribution. Oligocene : Florissant. 

Remarks. Cockerell maintains 'it appears to be allied to the ordinary species of 
Heriades . 

Heriades mersatus Cockerell 1923 

♦1923 Heriades mersatus Cockerell : 371. 
1 93 1 Heriades mersatus Cockerell ; Salt : 146. 

HoLOTYPE. Supposedly in Colorado Museum of Natural History, but not referred 
to by Rodeck (1938). 

Distribution. Oligocene : Florissant. 

Remarks. Nearest to H. halictimts. Author 'sure' of genus. 

Heriades mildredae Cockerell 1925 

♦1925a Heriades mildredae Cockerell : 420. 
193 1 Heriades mildredae Cockerell ; Salt : 146. 
1938 Heriades mildredae Cockerell ; Rodeck : 296. 



HoLOTYPE. Univ. of Colorado Mus., No. 15075, with counterpart. 

Distribution. Oligocene : Florissant. 

Remarks. Cockerell remarks that this species differs conspicuously from all 

other Heriades. 

Heriades priscus Cockerell 1917 

♦1917 Heriades priscus Cockerell : 383. 
1 93 1 Heriades priscus Cockerell ; Salt : 146. 

HoLOTYPE. U.S. Nat. Mus., No. 62543. 

Distribution. Oligocene : Florissant. 

Remarks. The largest Heriades from Florissant. A robust, hairy, black insect. 
Cockerell did not appear to have much doubt about its generic position. 

Heriades saxosus Cockerell 1913 

♦1913b Heriades saxosus Cockerell : 233. 
1931 Heriades saxosus Cockerell ; Salt : 146. 
1938 Heriades saxosus Cockerell ; Rodeck : 296. 

HoLOTYPE. Univ. of Colorado Mus., No. 18638. 
Distribution. Oligocene : Florissant. 
Remarks. Nearest to H. laminarum Cockerell. 

Genus MEGACHILE LatreiUe 1802 

* 1 802a Megachile Latreille : 434. 
1810 Megachile Latreille ; Latreille : 439. 
1828 Megachile Latreille ; Curtis : pi. 218. 
1841 Chalicodoma Lepeletier : 309. 

Genotype. Apis centuncularis Linnaeus 1758, selected by Curtis (1828). 

Remarks. The International Commission on Zoological Nomenclature recom- 
mended that the type-fixation (i.e. Apis muraria Retzius 1783) of Latreille be set 
aside and that of Curtis (1828) upheld (ICZN Opinion 219, 1954). 

Megachile amagiiensis Cockerell 1925 

♦1925b Megachile amaguensis Cockerell : 7, pi. i, fig. i. 
1931 Megachile amaguensis Cockerell ; Salt : 146. 

HoLOTYPE. U.S. Nat. Mus., No. 69601. 

Distribution. Tertiary : Kudia River, Siberia. 

Remarks. Cockerell states, 'The distance of the first recurrent nervure from the 
base of the second submarginal cell is an unusual feature, which I do not observe in 
living species'. 


Megachile praedicta Cockerell 1908 

* 1 908c Megachile praedicta Cockerell : 31. 
1 93 1 Megachile praedicta Cockerell ; Salt : 146. 

HoLOTYPE. Cockerell Collection, whereabouts unknown. 

Distribution. Oligocene : Florissant. 

Remarks. Genus confirmed by leaves cut through as though by Megachile. 

Megachile sp. indet. (Brischke) 

1886 Chalicodoma Brischke : 278. 

1 89 1 Chalicodoma Brischke ; Scudder : 694 (5985). 

1907 Chalicodoma Brischke ; Handlirsch : 888. 

1931 Chalicodoma sp. Brischke ; Salt : 144. 

Distribution. Eocene : Baltic amber. 

Remarks. Chalicodoma Lepeletier = Megachile Latreille. An unnamed Chali- 
codoma was said by Brischke to occur in the Baltic amber, two specimens apparently 
being known to him. 

Genus OS MIA Panzer 1806 

*i8o6 Osmia Panzer : 230. 
1 8 10 Osmia Panzer ; Latreille : 439. 

Genotype. Apis bicornis Linnaeus 1758, = Apis rufa Linnaeus 1758, selected 
by Latreille (1810 : 439). 

Remarks. It has been found difficult to determine the identity of the species of 
Osmia known to Heer. The following notes may prove useful in clarifying this point. 

In 1865 Heer mentioned 'Drei Osmien', having previously (1849) described one 
specimen, Osmia antiqua, in the Karlsruhe Collection. But quite a number of so- 
called Osmia are now in the collection at Zurich. These possess Heer's manuscript 
names and were actually placed by him in four groups : (i) Osmia primaeva ; (2) 
Osmia sp. ? dubia ; (3) Osmia immortua and (4) Osmia kirbyana. Which of these 
were the three referred to in 1865 is not known. 

Cockerell (1909) studied the species 0. primaeva (see p. 167) and referred it to 
Andrena. He did the same for 0. kirbyana (see p. 202) and considered it, probably 
rightly, to be a wasp. No more 'Osmia examined by Heer were studied by Cockerell, 
except a now unidentifiable insect 9-5 mm in length. 

Of the remaining two groups, those specimens labelled as 0. immortua are un- 
fortunately indeterminable, while of the two specimens labelled 'dubia' , one is 
superficially much like immortua and being also indeterminable it is best placed 
there (p. 187). The second specimen, however, is good and has been selected as the 
type of a new species (p. 185). However, 'dubia , apart from being a manuscript 
descriptive term and not a name, is preoccupied by Osmia dubia (Germar). We 
have, therefore, named it 0. nigra. [See also Appendix, p. 256.] 


Osmia carbonum Heyden 1862 

*i862 Osmia carbonum Heyden : 75, pi. 10, figs 11, 12. 

1 89 1 Osmia carbonum Heyden ; Scudder : 721 (6219). 

1907 'Osmia carbonum' Heyden ; Cockerell : 228. 

1907 Osmia carbonum Heyden ; Handlirsch : 889. 

1936 Osmia carbonum Heyden ; Statz : 260. 

HoLOTYPE. British Museum (Natural History), 58779 (Heyden's 1862 fig. 11). 
Paratype. British Museum (Natural History), 58768 (Heyden's 1862 fig. 12). 
Other specimens. Mus. Comp. Zool. Harvard. 
Distribution. Oligocene : Rott. 

Remarks. Heyden's originals were badly figured. His fig. 11 is of an insect 
without visible wings, and the original of fig. 12, of which the counterpart existed, 
was a somewhat smaller insect which has at least the more proximal portions of the 
wings still preserved. Both are in the British Museum (Natural History) Collection, 
but were badly damaged by exposure to water during the war so that no information 
additional to that given by Heyden can be derived from them. 

On the other hand, the specimens studied by Cockerell (1907) at Harvard were 'a 
very good specimen with reverse' and another apparently unnamed one which may 
be the counterpart of the paratype. Cockerell says, 'it is evident that they were not 
part of Heyden's material as they do not agree with his figures and descriptions'. 
This is certainly true as regards his 'good specimen with reverse', but his statement 
relating to wingless originals of 0. carbonum is not quite correct as far as the paratype 
is concerned.^ Moreover, after mentioning the 'goodness' of the specimen with 
counterpart at Harvard, he later quite casually states it 'is indeterminate', a state- 
ment difficult to reconcile with others previously made. It now remains to be seen 
whether Cockerell's second specimen is the counterpart of Heyden's paratype. 
Should this prove to be the case, no change of nomenclature will be necessary. On 
the other hand, a significant difference will necessitate placing 0. carbonum under 
the gen. indet. group, and a new name will be required for Cockerell's specimens. 

Osmia antiqua Heer 1849 

*i849 Osmia antiqua Heer : 95, pi. 7, figs 2a, b. 

1856 Osmia antiqua Heer ; Giebel : 182. 

1865 Drei Osmien ; Heer : 386 {partim). 

1876 Osmiae ; Heer : 43 [partim). 

1891 Osmia antiqua Heer ; Scudder : 721 (6218). 

1907 Osmia antiqua Heer ; Handlirsch : 889. 

1909c 'Osmia' Cockerell : 316. 

1938 Osmia Armbruster : 87. 

3 We consider it quite possible that the specimens described by Cockerell (1907) are, in part, the same 
as those described by Heyden (1862) from the collection of Dr Krantz. The difficulty of identification 
almost certainly lies with Heyden's poor descriptions and illustrations. That certain irreconcilable 
anomalies, however, do exist is quite clear, e.g. the statement that no signs of visible wings are present 
in a specimen of Heyden's Apis dormitans whereas all Cockerell's specimens of Apis (from Rott) possess 
them. In the circumstances, we have decided either to be cautious where actual similarities suggest 
themselves, as in Osmia carbonum, or to regard the two sets of material as separate and distinct, as for 
Apis dormitans. 


HoLOTYPE. Badische Landessammlung fiir Naturkunde, Karlsruhe (Oe.91), 

Distribution. Miocene : Oeningen. 

Remarks. Cockerell (1909c) said this 'cannot apparently be referred to Osmia 
or any other genus with certainty'. He, however, never saw the type and is beheved 
to have based his statement on Heer's (1849) entry of a specimen from the Karlsruhe 
Collection, and his rather poor illustration of it. Whether Heer (1865) intended this 
specimen to be included in his 'Drei Osmien' is not known, but it is assumed to be so. 

We have now studied the type and, although not well preserved, the following 
characters can be discerned. Broad head with a suggestion of large compound eyes ; 
parts of one fore and two hind legs, the latter, on the right, possessing distinguishable 
hairs ; thorax well chitinized, probably not so broad as head ; abdomen also well 
chitinized, some segmentation visible, quite broad ; some details of wing venation 
can be discerned, but since hind- and forewing overlap, and parts are missing, it is 
only possible to state that what can be seen is not incompatible with the wing 
venation of Osmia, and certainly the character of the hind legs, segmentation of 
abdomen, colour, general habitus etc. do not rule out Osmia either. 

Measurements. Overall length, 12 mm ; breadth of head, 4-5 mm (approx.) ; 
breadth of abdomen 5 mm ; length of abdomen, 7 mm ; length of forewing, 9 mm. 

Osmia nigra sp. nov. 
(PI. I, figs 5, 6) 
HoLOTYPE. Eidgenossische Technische Hochschule, Zurich, PI. I, 719. 
Paratype. Same collection, PI. I, 685. 
Distribution. Miocene : Oeningen. 

Description and Remarks. The holotype (labelled Osmia sp. ? duhia) is a 
comparatively small, black, insect, which is preserved in two parts, the abdomien 
being 5 mm distant from the head and thorax, but evidently belonging to it. The 
head is very imperfectly preserved but is as broad as the thorax. The thorax 
shows a well-developed mesoscutum. Parts of the middle and the two hind legs 
are preserved, and what is believed to be the hind tibia is triangular-shaped. The 
hind femur, however, is as strong and almost as broad as the tibia. Only the left 
forewing is preserved, but most of the venation can be discerned and it agrees well 
with Osmia : short, tapering marginal cell, broader at the base where it joins the 
pterostigma about the middle ; pterostigma well defined but not particularly large ; 
radial vein very strong ; two subequal submarginal cells ; basal vein short and 
relatively straight, of two almost equal segments, only slightly deflected at their 
junction ; cross-vein cu-a coincident with that of basal vein at point of juncture with 
Cu, but posteriorly it arches slightly proximad. Abdomen clearly reveals inter- 
segmental membranes in the distal parts. 

The paratype is an insect very reminiscent of the holotype and can be regarded 
as conspecific. The thorax is well developed with a broad convex mesoscutum. A 


middle and two hind legs are apparently preserved, with triangular tibiae. Some 
aspects of the abdomen are clearly shown. The specimen, however, is not generally 
well preserved but nevertheless corresponds with the holotype in all those characters 
that can be discerned except for a slightly lighter coloration. Since darker patches 
exist on various parts of the body, however, it is not known which was the more 
natural coloration for this species. No wings are preserved. Heer's manuscript 
name for this specimen was Apis adamitica, but it was clearly not the type of the 
latter species. 

Measurements. Holotype : length of abdomen, 6-5 mm ; width, 4-5 mm ; 
forewing length 5-5 mm (approx.). Paratype : length of abdomen 6 mm ; length 
of thorax and abdomen (together) 10 mm. 

Osmia sp. indet. (Menge) 

1856 Osmia Menge : 26. 

1891 05mm Menge ; Scudder : 721 (6217). 

1907 Osmia Menge ; Handlirsch : 889. 

1931 Osmia sp. Menge ; Salt : 144. 

Distribution. Eocene : Baltic amber. 

Remarks. Specimen referred to but neither described nor species determined. 

Family MEGAGHILIDAE incertae sedis 
*Apiaria' dubia Germar 1849 

♦1849 Apiaria dubia Germar : 66, pi. 2, fig. 8. 

1856 Osmia dubia Germar ; Giebel : 182. 

1891 Apiaria dubia Germar ; Scudder : 688 (5935). 

1 89 1 Osmia dubia Giebel ; Scudder : 721 (6220). 

1907 (Osmia) dubia Germar ; Handlirsch : 889. 

Distribution. Oligocene : Orsberg bei Bonn. 

Remarks. Nothing appears to be known about this specimen. The correctness 
of Giebel's generic determination is very doubtful. 

Megachilidae gen. et sp. indet. 

Material. Eidgenossische Technische Hochschule, Zurich, Nos. PI. I, 687 (with 
counterpart) ; PI. I, 688 (with counterpart PL I, 689) ; X ; and PL I, 718. 

Distribution. Miocene : Oeningen. 

Description and Remarks. Two of the specimens bear Heer's MS label 
'immortua'. The specimens appear to be bees, whose known parts are not incom- 
patible with Osmia but whose real systematic position cannot be satisfactorily 
ascertained owing to lack of diagnostic characters. The state of preservation is as 


(i). PI. I, 687 (and counterpart). A fairly well-chitinized insect with head, 
thorax, abdomen and portion of one antenna preserved, but with almost all traces 
of wings effaced. Body length 10 mm ; length of abdomen 5-5 mm, width 3 mm ; 
length of thorax 3-25 mm, width 3 mm. Apart from segments of the abdomen few 
morphological details can be discerned. 

(2). PI. I, 688 and counterpart PI. I, 689. A poorly-defined specimen, no longer 
with visible wing venation though the outline of the wings themselves can be detected. 
Tegulae quite prominent. Body length 11 mm approx. ; length of abdomen 
5*5 mm, width 375 mm ; length of thorax not accurately determinable, width 
3-25 mm. 

(3). One specimen marked 'X, Hymenoptera dub.' Body chitinized, dark brown 
in colour and comparatively well preserved, but legs and wings completely lacking. 
Two well-developed compound eyes present ; thorax, though less well outlined, is 
featureless ; abdominal segments, five in number, clearly in evidence. Body length 
10-25 mm ; length of abdomen 5 mm, width 3-25 mm ; length of thorax 3-25 mm, 
width 3 mm. 

(4). PI. I, 718, bearing Heer's MS label Osmia sp. ? {dubia). Head, thorax and 
abdomen are preserved, but legs are lacking and the wing venation is too indistinct 
for satisfactory recognition. Both the head and thorax are well chitinized and dark 
brown in colour, but the abdomen is paler. Body length (allowing for curved 
position) 10 mm approx. ; length of abdomen 5 mm, width 3-5 mm ; length of 
thorax 3-1 mm, width 3 mm (approx.). 

Family ANTHOPHORIDAE Dahlbom 1835 

Diagnosis. Labrum usually broader than long, attached by rather short margin 
to clypeus ; subantennal areas absent ; subantennal sutures directed towards inner 
margins of antennal sockets ; lower sides of clypeus frequently bent back parallel 
to long axis of body ; facial foveae almost always absent ; galeae elongate post- 
palpally, short pre-palpally ; labial palpi with first two segments elongate and 
flattened, first segment at least as long as second ; glossa linear, usually with a 
flabellum ; pre-episternal suture usually absent below scrobal suture, present above 
it ; middle coxae elongate, except in a few parasitic forms ; basitibial plates usually 
present ; scopa, when present, occurs on the posterior tibiae and basitarsi, and very 
rarely found on abdominal sternites as well ; pygidial area often present. 

Usually three submarginal cells ; when two only, second often much shorter than 
first ; rarely only one closed submarginal cell. 

Distribution. Eocene to Recent. 

Subfamily XYLOCOPINAE Latreille 1802 

Diagnosis. Clypeus not protuberant but relatively flat, lateral portions seen 
from beneath a little bent backward and more nearly transverse than longitudinal ; 
pygidial plate absent, represented in some females by a flat-topped apical spine 
which, unfortunately, is hidden in a dense pygidial fimbria and hence would be 


difficult to see in fossils ; scopa of female not forming a corbicula ; inner apical 
margins of posterior tibiae bare or hairy, without a hind tibial rake. 

Michener (1944) also mentions a further distinctive feature of the clypeus of 
Xylocopinae, though of doubtful use for fossils. The portions of the epistomal 
suture between the anterior tentorial pits and the dorsolateral angles of the clypeus 
are subparallel to one another, while below the pits the clypeus broadens abruptly. 

Remarks. This subfamily contains two distinctive tribes, the Ceratinini and the 
Xylocopini, both represented by fossil forms. 

Tribe CERATININI Latreille 1802 

Diagnosis. Small, rather slender bees with little pubescence ; mandibles very 
broad basally but abruptly narrowing to the slender, subparallel-sided apical 
portions ; metanotum horizontal ; apex of seventh tergum of the female produced 
to a small, sharp, median point. 

Wing surfaces hairy throughout, not papillate ; marginal cell broad, longer than 
distance from its own apex to wing tip, and considerably bent from the wing margin 
apically ; jugal lobe of each hindwing is one-third to half as long as the anal lobe. 

Remarks. Only one fossil specimen so far known. 

Genus CERATINA Latreille 1802 

1802a Clavicera Latreille : 432. 
*i8o2b Ceratina Latreille : 380. 

Genotype. [Hylaeus alhilahris Fabricius 1793) = Apis cucurhitina Rossi 1792, 
by monotypy. 

Remarks. A recommendation is before the International Commission on Zoo- 
logical Nomenclature that the name Clavicera be suppressed. 

Ceratina disrupta Cockerell 1906 

♦1906 Ceratina disrupta Cockerell -.37. 
1907 Ceratina Cockerell ; Handlirsch : 891. 
1973 Ceratina disrupta Cockerell ; Daly : 15, pi. 2, text-fig. 4b. 

HoLOTYPE. Mus. Comp. Zool. Harvard, No. 2001 (= Scudder Coll. No. 9355). 
Distribution. Oligocene : Florissant. 

Remarks. This black, average-sized bee (8 mm) has the general appearance of 
a Ceratina. Description of forewing given by Cockerell. 

Tribe XYLOCOPINI Latreille 1802 

Diagnosis. Large, robust bees with coarse, plumose hair ; metanotum vertical ; 
apex of seventh tergum of female produced to a flat-topped spine, largely hidden in a 
dense mass of hair ; basitarsi fully as long as the corresponding tibiae. 


Wings elongate, papillate distally ; marginal cell very slender, its apex bent away 
from the wing margin and the cell itself much longer than the distance from its own 
apex to the wing tip ; jugal lobe of hindwing much shorter than the anal lobe. 

Remarks. All fossils have been assigned to the genus Xylocopa. 

Genus XYLOCOPA Latreille 1802 

1802a Xilocopa Latreille : 432. 
* 1 802b Xylocopa Latreille : 379. 
1810 Xylocopa Latreille ; Latreille : 439. 

Genotype. Apis violacea Linnaeus 1758, selected by Latreille (1810 : 439). 

Remarks. The name Xylocopa has been placed on the list of nomina conservanda 
(ICZN Opinion 743). 

Xylocopa friesei Statz 1936 
♦1936 Xylocopa friesei Statz : 284, 293, pi. 8, fig. 37. 

HoLOTYPE. Statz Collection. 

Distribution. Oligocene : Rott. 

Remarks. Wing, thorax and hind basitarsus all strongly suggest a Xylocopa. 
According to Statz, the specimen is similar in size to Xylocopa senilis Heer, but since 
only the thorax, right forewing and four legs (two very fragmentary) are known, 
his statement can have but limited value. 

Xylocopa hydrobiae Zeuner 1938 
♦1938 Xylocopa hydrobiae Zeuner : 127, figs 20, 21. 

HoLOTYPE. Nat. Hist. Mus. Mainz, Hydrobienkalk Collection, No. 123. 

Paratype. Same collection, No. 5. 

Distribution. Miocene : Biebrich nr Mainz. 

Remarks. Vein 2nd r-m together with distal segment of Rs is Z-shaped. 

Xylocopa jurinei (Heer 1865) 

''1865 Bombus jurinei Heer : 386, fig. 296. 

1867 Bombus jurinei Heer ; Heer : 4, pi. 3, fig. 8. 

1872 Bombus jurinei Heer ; Heer : 473, fig. 296. 

1876 Bombus jurinei Heer : Heer : 4^. fis:. 206. 

1876 Bombus jurinei Heer ; Heer : 43, fig. 296. 

1879 Bombus jurinei Heer ; Heer : 411, fig. 338. 

1891 Bombus jurinei Heer ; Scudder : 690 (5953) 

1907 Bombus jurinei Heer ; Handlirsch : 892. 

1909c Xylocopa jurinei (Heer) ; Cockerell : 315. 

1928 Bombus jurinei Heer ; Wheeler : 97. 

1928 Xylocopa Wheeler : 97. 


1 93 1 Bombus jurinei Heer ; Cockerell : 301. 
1 93 1 Xylocopa Latreille ; Cockerell : 301. 
1938 Bombus jurinei Heer ; Armbruster : 87. 

HoLOTYPE. Eidgenossische Technische Hochschule, Zurich, No. PI. I, 682a 
(Heer 1865 : fig. 296). 
Other material. Same collection, No. PI. I, 682b (Heer 1867 : pi. 3, fig. 8). 
Distribution. Miocene (Sarmatian) : Oeningen. 

Remarks. If the specimen No. PI. I, 682b is not the counterpart of the holotype, 
a point very difficult to decide, then Cockerell (igogc) did not see the type, since his 
specimen was that figured by Heer (1867). 

Of this specimen, Cockerell (1909c) states, 'Only part of the venation can be made 
out, but all that can be seen agrees with Xylocopa, and not with Bombus'. Arm- 
bruster (1938) also suggests the name Xylocopa for this specimen. Having examined 
the two existing specimens, we endorse Cockerell's conclusion as applicable to both. 
Moreover, the tibia and basitarsus of the hind legs bear a well-developed scopa, which 
is typical of Xylocopa. Furthermore, in specimen PI. I, 682b, the basitarsus of the 
hind leg is approximately one and a half times as long as the tibia. This greater 
length of the basitarsus is a character of Xylocopa, whilst in Bombus the basitarsus 
is usually shorter than the tibia. The suture between mesoscutum and mesoscutel- 
lum is very distinct in the fossils and reminiscent of Xylocopa. A prominent feature 
of specimen PI. I, 682b is the strong and copious hairs protruding from the apex of the 
abdomen, which again is observed in some Xylocopa. 

The species is distinguished from Xylocopa senilis Heer by its larger size. 

Measurements. Holotype : length of thorax plus abdomen, 21 mm ; length of 
abdomen 14 mm, width 11 mm ; length of thorax 7 mm, width 8 mm ; length of 
fore wing 15 mm. 

Specimen PI. I, 682b : length of thorax plus abdomen 25 mm ; length of abdomen 
18 mm, width 11-5 mm ; length of thorax 7 mm, width 8 mm ; length of forewing 
16-5 mm. The legs here are well preserved, the length of the tibia being 375 mm 
and the length of the basitarsus 6 mm. It will also be noticed that the measurements 
of the thorax and the abdomen do not quite tally with those given by Heer. 

Both specimens have the head missing. 

Xylocopa senilis Heer 1849 
(PI. I, figs 3, 4) 

*i849 Xylocopa senilis Heer : 93, pi. 7, figs la, b, c. 

1856 Xylocopa senilis Heer ; Giebel : 181. 

1865 Xylocopa senilis Heer ; Heer : 386, fig. 295. 

1872 Xylocopa senilis Heer ; Heer : 473, fig. 295. 

1876 Xylocopa senilis Heer ; Heer : 43, fig. 295. 

1879 Xylocopa senilis Heer ; Heer : 411, fi.g. 337. 

1885 Xylocopa senilis Heer ; Scudder : 819, fig. 1109. 

1887 Xylocopa senilis Heer ; Scudder: 821, fig. 1126. 

1891 Xylocopa senilis Heer ; Scudder : 734 (6334). 

1907 Xylocopa senilis Heer ; Handlirsch : 891. 


1909c Xylocopa senilis Heer ; Cockerell : 415. 

1931 Xylocopa senilis Heer ; Pongracz : 117, pis (5) 7, (6) 8. 

1938 Xylocopa senilis Heer ; Armbruster : 88. 

HoLOTYPE. Badische Landessammlung fiir Naturkunde, Karlsruhe, Inv.1267. 
Paratype. Same collection. 

Other material. Same collection Inv.1270 (as photographed by Pongracz 
1931 : pi. (5) 7) ; Inv.1175 (Oe.89, no. 145) ; Inv.1175 (Oe.89, no. 146) ; Inv.1228 
(Oe.93). Also Eidgenossische Technische Hochschule, Zurich, no. 21a with counter- 
part no. 2ib, and another specimen doubtfully referred here, no. 21c. 

Distribution. Miocene : Oeningen. 

Remarks. We have studied specimens Inv.1175 (iio- 145)^ Inv.1175 (no. 146) 
and Inv.1228 from Karlsruhe, and specimens 21a, b and c from Zurich. It is quite 
clear that we have not seen the holotype in spite of the fact that we made a thorough 
search for it at Karlsruhe, but we believe a photograph of it was published by 
Pongracz (1931c : pi. (6) 8). The strongest evidence for this is the clear reference 
to the well-preserved antennae. The paratype, as defined by Heer, was 'ein sehr 
undeutliches Stiick aus derselben Sammlung'. That we have not seen this specimen 
is again evident. Possibly it is the second one photographed by Pongracz (1931 : 
pi. (5) 7) although the outline here is quite clear, and not indistinct as Heer suggested. 
Both types may now be lost but this cannot by any means be asserted since much 
material is still stored below the ruins of what was once the Badische Landessamm- 
lung, Karlsruhe. 

In the absence of types it is difficult to know whether this is a genuine Xylocopa 
or not. At least, the specimens at Karlsruhe that have been examined are known 
to lack decisive detail for generic determination. The following are the measure- 
ments and characters that can be more reliably ascertained. 

Inv.1175 (no. 145). Overall length 20 mm ; breadth of abdomen 8 mm ; breadth 
of thorax 5 mm ; length of wing on left 18 mm ; length of wing on right 14 mm. 
The impression of this insect is distinct and possibly a single leg is displayed on the 

Inv.1175 (no. 146). Overall length 21-25 mm ; breadth of abdomen 8 mm; 
breadth of thorax 5 mm ; length of wing on left 15 mm ; that on right 17 mm. This 
insect resembles no. 145, though it is not the counterpart. It is less distinctly 

Inv.1228. Overall length 19 mm ; length of abdomen 12 mm ; breadth of ab- 
domen 8 mm ; breadth of thorax 6 mm. This is a large brownish-black insect in 
which a few proximal veins of the wing can still be discerned, but not enough to 
serve any useful purpose. Very few other features are preserved. 

On the other hand, the Zurich Collection contains a better preserved specimen 
(no. 2ia, b) which shows enough of the venation of the forewing to identify it as a 
Xylocopa, the basal vein being short and the '2nd discoidal' cell (i.e. 2nd M) elongate. 
The copious hairs on the basitarsus of what appears to be the middle leg and the 
shortness of the remaining tarsal segments support this identification. Since the 
size agrees very well with that of Xylocopa senilis Heer, the specimen is assigned to 


this Species without reservation. It is the best preserved specimen available, and 
has the following measurements : overall length 22 mm ; length of abdomen 12 mm, 
width 8-9 mm ; length of thorax 7 mm, width 9 mm ; length of forewing 8 mm. 
The head is rather crushed and thus of little value for measurement since it is too 
much spread out. The broad thorax indicates a distinct transverse suture of the 

Specimen no. 21c is poorly preserved, without recognizable venation. The 
abdomen appears to be more slender than normal for the species. Its specific and 
even generic identity is open to grave doubt, though it was labelled 'Xylocopa senilis 
X Hr. (?)' in Heer's time. It is a smaller insect, possibly little more than 10 mm 
long, with a greater constriction between thorax and abdomen. 

Subfamily ANTHOPHORINAE Dahlbom 1835 

Diagnosis. Clypeus almost always protuberant so that lateral portions, seen 
from below, are bent posteriorly and lie parallel to the long axis of the body ; the 
margins of the clypeus generally converge upward to the dorsolateral angles ; 
pygidial plate present in nearly all females and most males ; scopa of female not 
forming a corbicula ; anterior coxae usually little broader than long ; inner apical 
margins of posterior tibiae bare or hairy, without a tibial rake. 

Remarks. This very large subfamily is believed by Michener (1944) to be a 
monophyletic unit. The characters, however, of parasitic forms, which have evolved 
along different lines, are sometimes difficult to reconcile with those of non-parasitic 
forms, and are not always compatible with the diagnostic characters given above. 
The tribes Eucerini, Anthophorini and Melectini are represented by fossil forms. 
[See also Appendix, p. 256.] 

Tribe EUCERINI Latreille 1802 

Diagnosis. Large, pollen-collecting bees ; clypeus of the male usually partly 
yellow or white ; antennae of males usually much elongated ; flagellar segment in 
the female and in some males nearly as long as the scape ; posterior legs of the 
male only rarely enlarged or modified ; arolia present ; scopa present ; males often 
possess distinct pygidial and basitibial plates. 

Wing pubescent throughout and only feebly papillate apically ; marginal cell as 
long as, or longer than, the distance from its own apex to the wing tip, its apex 
narrowly rounded and bent gradually away from the wing margin ; most (including 
all North American) genera with three submarginal cells, the first and third longer 
than the second ; second abscissa of vein M-Cu of the posterior wings usually nearly 
twice as long as the transverse or somewhat oblique cu-a (comparable with 
Emphorini) and the jugal lobe about half as long as the anal lobe ; pterostigma 
reduced, tapering little, if at all, beyond the base of vein r. 

Remarks. The long paraglossae in the Eucerini are of doubtful diagnostic value 
for fossil forms. This tribe is one of the largest of the Anthophorinae but is re- 
presented with reasonable certainty by only one fossil specimen {Tetralonia herlandi) . 


Another species, Eucera mortua, described by Meunier (1920) has now been placed 
in Anthidium, but it is clear that Meunier felt very sure of his genus. The specimen, 
therefore, should be re-examined. That this was not done by us was due to our 
being mislead by Statz's erroneous statements (see Anthidium mortuum (Meunier), 
p. 178). 

Genus TETRALONIA Spinola 1838 

1810 Macrocera Latreille {nee Meigen) : 339. 
*i838 Tetralonia Spinola : 538. 

Genotype. Eucera antennata Fabricius 1793 (= Apis malvae Rossi 1790) by 
subsequent designation of Sandhouse (1943 : 603). 

Tetralonia berlandi Theobald 1937 
*i937 Tetralonia berlandi Theobald : 131, pi. 2, fig. ii ; pi. 10, fig. i. 

HoLOTYPE. Museum at Marseille, Celas Collection, no. C.31. 

Distribution. Oligocene, Sannoisien du Gard, France. 

Remarks. This broad, black bee most closely approaches to the Recent T. 
duvaucelii Lepel., according to Theobald. It is 10 mm long, with the forewing 
measuring 6 mm in length. The specimen is adequately described and figured, and 
Theobald's figure certainly suggests an anthophorid. 

Tribe ANTHOPHORINI Dahlbom 1835 

Diagnosis. Moderate-sized to large, pollen-collecting Anthophorinae ; clypeus 
of male usually pale ; posterior angles of mandibles are beneath posterior margins 
of eyes ; first flagellar segment often elongated, sometimes as long as the scape ; 
scopa not large, consisting of simple, or slightly plumose, hairs, or in some cases 
with some plumose hairs intermixed ; claws of male cleft, of female with median 
tooth on inner side ; arolia mostly present ; eighth tergum of male is usually bilobed, 
without a pygidial plate. 

Wings largely bare and strongly papillate apically ; pterostigma small, usually 
one and a half times as long as broad and ending at the base of vein r ; marginal cell 
slightly shorter than the distance from its own apex to the wing tip, rounded at its 
apex ; first submarginal cell, usually slightly longer on posterior margin than are the 
subequal second and third cells ; jugal lobe of posterior wing usually a little over 
one-third as long as anal lobe ; second abscissa of vein M-Cu of posterior wings about 
as long as cu-a. 

Remarks. The genera Anthophora and Anthophorites are represented by fossil 
forms. The latter genus was erected by Heer (1849) for all fossil bees with affinities 
with Anthophora. The type, Anthophorites mellona Heer, therefore, is in need of 
further examination to establish its true relationship. 


Genus ANTHOPHORA Latreille 1803 

1802a Podalirius Latreille : 430. 
*i8o3 Anthophora Latreille : 167. 
1810 Anthophora Latreille ; Latreille : 439. 

Genotype. Megilla pilipes Fabricius 1775 (= Apis acervorum Linnaeus 1758), 
selected by Latreille (1810). 

Remarks. The International Commission of Zoological Nomemclature has 
recommended that the generic name Podalirius Latreille (1802) be rejected (ICZN 
Opinion 151, 1944). 

Anthophora melfordi Cockerell 1908 

(PL I, fig. I) 

♦1908b Anthophora melfordi Cockerell : 323. 

i9o8d Anthophora melfordi Cockerell ; Cockerell : 575, fig. 6. 

1 93 1 Anthophora melfordi Cockerell ; Salt : 145. 

1938 Anthophora melfordi Cockerell ; Rodeck : 295. 

HoLOTYPE. British Museum (Natural History), 1. 9259. Counterpart in Colorado 
University Museum, No. 18922. 

Distribution. Oligocene : Florissant. 

Remarks. According to Cockerell (1908b), This is the first genuine fossil 
Anthophora' . The parts preserved include the head, thorax, first abdominal 
segment, parts of three legs, most of the forewing overlying remnants of the hind- 
wing and the extended mouth parts. 

The specimen has been re-examined and the following points noted. The fore- 
wing is well preserved, except distally. In the hindwing, however, since it is overlain 
by the forewing, the venation is less readily ascertainable. Some veins appear to 
be very badly preserved or not at all. The legs, although undoubtedly hair}/^, are 
possibly not so to the extent Cockerell suggests, nor are we able to verify his state- 
ment of the 'copious' quantity of hair on the 'tongue'. 

As regards the venation of the forewing, there is no need for us to differ from 
Cockerell's decision to place the specimen in Anthophora. The juncture, however, 
where the first recurrent vein reaches the second submarginal cell, may be 'near the 
beginning of its last third instead of at the middle', but it must also be mentioned 
that this juncture is far from being clearly visible. Vein cu-a of the hindwing is 
very oblique, as Cockerell states. 

Genus ANTHOPHORITES Heer 1849 

♦1849 Anthophorites Heer : 97. 
1909c Anthophorites Heer ; Cockerell : 315. 

Genotype. Anthophorites mellona Heer (1849), selected by Cockerell (1909c). 
Remarks. According to Cockerell, 'Anthophorites cannot be precisely defined, 
but includes various fossil bees supposed to be more or less similar to Anthophora' . 


The genus is obviously a repository for all Anthophorid-like insects that cannot 
otherwise be satisfactorily determined. 

The holotype of the genotypic species is now lost, but it was illustrated by Heer 
with some wing venation and body characters, insufficient for the establishment of a 
generic diagnosis. We have, however, refrained from sinking this genus because no 
specimens attributable to the type species have been examined by us. 

Anthophorites gaudryi Oustalet 1870 

♦1870 Anthophorites gaudryi Oustalet : 104, pi. 2, figs 11, 13. 

1891 Anthophorites gaudryi Oustalet ; Scudder : 687 (5922). 

1907 Anthophorites gaudryi Oustalet ; Cockerell : 227. 

1907 Anthophorites gaudryi Oustalet ; Handlirsch : 890. 

1937 Anthophorites gaudryi Oustalet ; Theobald : 433. 

Holotype, Museum at Corent, France. 

Distribution. Oligocene (Stampian) : Corent, France. 

Remarks. The specimen is poorly preserved and lacks the head. Scudder 
(1891) compared it with A. parietaria. Cockerell (1907) writes, 'has a curiously 
fly-like appearance', and 'its hairs are not plumose'. Oustalet's figure (1870 : pi. 2, 
fig. 11) is most difficult to interpret. 

It should be noted that the species is not listed by Theobald (1937 : 375) where 
A . mellona is given as occurring at Corent instead. 

Anthophorites thoracica Heer 1867 

♦1867 Anthophorites thoracica Heer : 6, pi. 3, fig. 14. 
1891 Anthophorites thoracicus Heer ; Scudder : 687 (5925). 
1907 Anthophorites thoracicus Heer ; Handlirsch : 890. 

Holotype. Possibly Badische Landessammlungen fiir Naturkunde, Karlsruhe. 

Distribution. Miocene : Radoboj. 

Remarks. Apparently based on one specimen from Radoboj. We particularly 
searched for this specimen at Karlsruhe without avail. It may nevertheless still 
reappear when the material now stored in the cellars there is unpacked and sorted. 

Anthophorites longaeva Heer 1867 

♦1867 Anthophorites longaeva Heer : 5, pi. 3, figs 12, 13. 

1 89 1 Anthophorites longaevus Heer ; Scudder : 687 (5923). 

1907 Anthophorites longaevus Heer ; Handlirsch : 890. 

1909c Anthophorites longaeva Heer ; Cockerell : 315. 

1938 Anthophorites longaeva Heer ; Armbruster : 87. 

Holotype. Heer's (1867) fig. 13 from Radoboj, Whereabouts unknown. 
Paratype. Heer's (1867) fig. 12 from Oeningen. Whereabouts unknown. 
Other Specimens. Eidgenossische Technische Hochschule, Zurich, Nos PL I, 
683, PL I, 712 and PL I, 809. 


Distribution. Lower and Upper Miocene : Radoboj and Oeningen, 

Remarks. Of the two specimens figured by Heer (1867), fig. 13, from Radoboj, 
appears to be well preserved. Fig. 12, from Oeningen, however, is described as 
'sehr ahnlich A. mellona, aber betrachtlich grosser'. It is believed to have been 
seen by Cockerell (1909c) who described it as a bee with large eyes, narrow face, 
middle joints of a rather stout flagellum a little longer than broad, and an abdomen 
broad at the base, with fine hair ; hind tibia 3-75 mm, hind basitarsus 2-5 mm and 
broadened ; venation indistinguishable. The generic position of the specimen was 
considered by Cockerell to be 'wholly obscure'. The three Zurich specimens can be 
described as follows. 

PI. I, 683. A strongly chitinized insect having the head partly lacking ; thorax 
well outlined, but without distinct diagnostic characters ; abdomen with segments 
clearly indicated and covered with hairs ; hind legs poorly preserved, the basitarsus 
and tibia apparently hairy. Body length 16 mm ; length of abdomen 10 mm, 
width 5 mm ; length of thorax 4 mm, width 4 mm. 

PL I, 712. Head, thorax and abdomen preserved, but not so highly chitinized as 
PI. I, 683 ; the strong costa of the wing can be discerned ; abdomen somewhat 
distended ; a triangular-shaped chitinous area that is preserved may be a tibia. 
Length of body 17 mm ; length of head 2-25 mm ; length of thorax 4-5 mm, width 
4-5 mm ; length of abdomen 10-25 ^^^^, width 575 mm. 

PL I, 809. A very weakly chitinized specimen on which few details can be dis- 
cerned. Head, thorax and abdomen present, wings completely lacking and legs 
practically absent. Length of body 17-5 mm ; length of head 2 mm ; length of 
thorax 5 mm, width 5 mm ; length of abdomen 10-5 mm, width 6 mm. 

These three specimens are not determinable. They are placed under this species 
because the characteristics that can be discerned are compatible with the holotype. 
They are labelled with the name A . longaeva in a nineteenth-century handwriting. 

Anthophorites mellona Heer 1849 

♦1849 Anthophorites mellona Heer : 97, pi. 7, figs 4a-c. 

1856 Anthophorites mellona Heer ; Giebel : 183. 

1891 Anthophorites mellona Heer ; Scudder : 687 (5924). 

1907 Anthophorites mellona Heer ; Handlirsch : 891. 

1907 Anthophorites mellona Heer ; Cockerell : 229. 

1909c Anthophorites mellona Heer ; Cockerell : 315. 

1914 Anthophorites mellona Heer ; Meunier : 193, pi. 6, figs 5a, b, text-fig. 5. 

1937 Anthophorites melona (Heer) ; Theobald : 375. 

Holotype. Originally in Eidgenossische Technische Hochschule, Zurich, but 
cannot now be traced. 

Other specimens. One specimen in Mus. Comp. Zool. Harvard, and another, 
described by Meunier, is believed to be in the Marseille Museum. 

Distribution. Miocene : Oeningen. Also reported from the Oligocene of Aix 
and the Oligocene of Cereste, Corent, Celas, France. 


Remarks. Heer's illustration is unsatisfactory. The thorax is very hairy, and 
some wing venation can be made out. One specimen only was described by Heer, 
from the Zurich collection, where however we have not succeeded in finding it. 

Cockerell (1907), while investigating specimens in the Museum of Comparative 
Zoology, Harvard, came across an example of Anthophorites mellona Heer, from 
Oeningen, determined by Heer himself. Its venation could not be discerned. Since, 
however, some wing venation must have existed in Heer's original, this specimen 
cannot be the holotype. Cockerell (1909c) fixed Anthophorites mellona as the 

Meunier's (1914) specimen from Aix-en-Provence was thought by him to belong 
to this species, though his description was meagre. He did, however, give photo- 
graphs of two specimens. His pi. 6, fig. 5a is the specimen described ; it appears 
to be well preserved. His pi. 6, fig. 5b is probably a photograph of Heer's A. 
mellona used for comparison, although no mention of this is given. Moreover, the 
resemblance between the two specimens is probably merely superficial. 

The reports from the Oligocene are even more doubtful. They appear in Theo- 
bald's (1937) table, for each of the three localities. There is only a single specimen, 
however, and one Anthophorites is indeed known from the Oligocene of Corent, 
France, viz. A. gaudryi (p. 195). It is listed by Theobald under that fauna (1937 : 
433), but omitted from the list. 

Anthophorites titania Heer 1849 

*i849 Anthophorites titania Heer : 99, pi. 7, figs 5a, b. 

1856 Anthophorites titania Heer ; Giebel : 183. 

1891 Anthophorites titania Heer ; Scudder : 687 (5926). 

1907 Anthophorites titania Heer ; Handlirsch : 891. 

1909c Anthophorites titania (Heer) ; Cockerell : 315. 

Holotype. Badische Landessammlung fur Naturkunde, Karlsruhe, Inv.1273 

Paratype. Eidgenossische Technische Hochschule, Zurich, No. PL I, 686 (not 
figured by Heer). 

Distribution. Miocene : Oeningen. 

Description. The holotype has been examined, and found to have the head 
missing ; the thorax is much distorted, and somewhat hairy. Both hind legs are 
in part preserved, one tibia and two basitarsi being very distinct. Both tibiae and 
basitarsi are broad and very hairy. The tibia is less well preserved but appears to 
be more than one and a half times as long as the basitarsus, without however being 
much broader. The abdominal segments are clearly visible and covered with long 
hairs. The basal vein is strongly curved at its base and soon straightens out. 
Length of abdomen 9 mm, width 5-5 mm. 

The paratype, stated by Heer (1849) to have once been in the Karlsruhe Collection, 
is now at Zurich and has been examined by us. It was also studied by Cockerell 
(1909c) who considered its generic position to be obscure. This we hereby endorse. 


The specimen is without head or legs, but the segmentation of the abdomen is clear. 
A few details of the thorax can be discerned : the pronotum is narrow, well margined 
but not quite reaching the tegulae ; the mesonotum is prominent and arched, but 
rather small and well defined posteriorly by a transcutal suture ; the mesoscutellum 
is clearly evident, the metathorax is delimited posteriorly by 'a sharp edge separating 
base from apical truncation', and a definite constriction exists between it and the 
abdomen. The wing venation has been almost effaced, but a distinct pterostigma 
still remains and a strong radius is present. Length of body (without head) 
13-25 mm ; length of thorax 4-25 mm, width 4-1 mm ; length of abdomen 9 mm, 
width 5-25 mm ; length of forewing 8 mm (as near as can be ascertained). 

Remarks. The shape of the basal vein of the holotype is a trait not uncommon 
in Anthophora, as the examination of many Recent specimens has revealed. It 
argues strongly against the alternative of assigning A . titania to Halictus. But the 
characters preserved do not permit us to be certain that the species is an Anthophora 
sensu stricto. 

Anthophorites tonsa Heer 1849 

♦1849 Anthophorites tonsa Heer : 99, pi. 7, figs 6a, b. 

1856 Anthophorites tonsa Heer ; Giebel : 183-184. 

1 891 Anthophorites tonsus Heer ; Scudder : 687 (5927). 

1907 Anthophorites tonsus Heer ; Handlirsch : 891. 

Holotype. Badische Landessammlung fiir Naturkunde, Karlsruhe. 

Distribution. Miocene : Oeningen. 

Remarks. One specimen only has been described, and it cannot at the moment 
be found at Karlsruhe because of war damage. Heer's (1849) illustration is un- 

Anthophorites veterana Heer 1849 

*i849 Anthophorites veterana Heer : 100, pi. 7, figs 7a, b. 

1856 Anthophorites veterana Heer ; Giebel : 183-184. 

1891 Anthophorites veteranus Heer ; Scudder : 687 (5928). 

1907 Anthophorites veteranus Heer ; Handlirsch : 891. 

Holotype. Eidgenossische Technische Hochschule (Lavater Collection), Zurich, 
No. 14. 

Other specimen. Same collection, No. PL I, 684. 

Distribution. Miocene : Oeningen. 

Description and Remarks. Heer's figure of the holotype agrees well with the 
holotype itself except that he omits the legs although he mentions them in the text. 
It is not well preserved. The insect was robust, with a broad convex mesoscutum. 
Tegulae prominent. The hind leg, particularly the basitarsus, is hairy and as broad 
as the tibia. On account of a depression in the rock which crosses the tibia, the 
relative lengths of the tibia and basitarsus are difficult to assess. Some venation 


is present, but since the forewing overlies the hindwing, it is impossible to decipher 
it satisfactorily. The strong radius, however, is distinct and also some proximal 
portions of M + Cu and Cu, Length of body 14 mm ; length of thorax 4-25 mm, 
width 5-5 mm ; length of abdomen 875 mm, width 6-5 mm ; forewing length 11 mm. 
A second specimen (PI. I, 684) labelled Anthophorites may tentatively be placed 
here. It is 12-5 mm long, extremely poorly preserved and black in colour. 

Tribe MELECTINI Westwood 1840 

Diagnosis. Large, parasitic bees ; clypeus of male black ; posterior angle of 
mandible lies beneath middle of lower end of eye ; first flagellar segment not 
unusually elongated ; scopa absent ; basitibial plates absent ; claws with large 
inner basal lobe or tooth ; middle tibial spurs unmodified ; eighth tergum of the 
male usually feebly bilobate or bispinose and usually without a pygidial plate ; 
seventh abdominal tergite of female with pygidial plate ; seventh sternite of female 
somewhat produced, with the margins upturned to form a semitubular guide for 
the sting. 

Pterostigma several times as long as broad, narrow, and not tapering beyond the 
base of vein r ; marginal cell shorter than the distance from its own apex to the wing 
tip, broadly rounded apically, and usually not protruding beyond the last submarginal 
cell ; normally three submarginal cells, the first usually longer than either of the 
others ; second abscissa of vein M-Cu of posterior wings shorter than vein cu-a, 
sometimes virtually absent ; jugal lobe of hind wing very small. 

Genus PROTOMELECTA Cockerell 1908 

*i9o8a Protomelecta Cockerell : 341. 
1909b Protomelecta Cockerell ; Cockerell : 81. 

Genotype. Protomelecta brevipennis Cockerell by monotypj^ 

Remarks. According to Cockerell, 'A very Meleda-like genus but with the 

venation of the Eucerines and the large stigma of the Andrenids'. Special mention 

of the type was made by Cockerell (1909b). 

Protomelecta brevipennis Cockerell 1908 

* 1908a Protomelecta brevipennis Cockerell : 341. 
1909b Protomelecta brevipennis ; Cockerell : 81. 
1938 Protomelecta brevipennis Cockerell ; Rodeck : 298. 

HoLOTYPE. Univ. Colorado Mus., No. 18639. 

Distribution. Oligocene : Florissant. 

Remarks. The affinities of this species are by no means clear. It would appear 
to be an anthophorid, but its relationship with the melectines depends to some 
considerable extent on theoretical considerations only. A reinvestigation of the 
specimen is most desirable. 


Family ANTHOPHORIDAE incertae sedis 

* Anthophora' effossa Heyden 1862 

♦1862 Anthophora effossa Heyden : 76, pi. 10, fig. 10. 

1891 Anthophora effossa Heyden ; Scudder : 687 (5921). 

1907 Anthophora effossa Heyden ; Handlirsch : 890. 

1907 Anthophora effossa Heyden ; Cockerell : 228. 

1936 Anthophora effossa Heyden ; Statz : 260. 

HoLOTYPE. British Museum (Natural History), 58781. 

Distribution. Oligocene : Rott. 

Remarks. It is necessary to distinguish between the 'Anthophora effossa' of 
Cockerell (1907) which appears to be an ant (p. 202), and the Anthophora effossa 
Heyden which Cockerell apparently did not see. He maintained that Heyden's 
figure and description did not agree with his specimen, though whether he was 
sufficiently indulgent towards Heyden's work in this respect is not clear. Heyden's 
figures, however, are entirely useless for diagnostic purposes, and the holotype was 
damaged during the war, so that details are no longer recognizable. A further point 
to be noted is the very arbitrary way in which Heyden referred his specimen to 
Anthophora. For these reasons one is left with no alternative but to place it incertae 

Anthophoridae gen. et sp. indet. i 

1856 Anthophora Latreille ; Menge : 26. 

1886 Anthophora ? Latreille ; Brischke : 278. 

1 89 1 Anthophora ? Latreille ; Scudder : 686 (5919). 

1 89 1 Anthophora } l.d±xei\\Q ; Scudder: 686(5920). 

1907 Anthophora ? Latreille ; Handlirsch : 890. 

Distribution. Eocene : Baltic amber. 

Remarks. Since Brischke had access to, and reported on, Menge's specimens, 
it is quite possible that Brischke refers here to Menge's Anthophora. Four specimens 
appear to have existed, though Menge mentioned only one. 

Anthophoridae gen. et sp. indet. 2 

1831 Hymenopt. aus der Fam. der Bienen ; Hiinefeld : 2000 (i.e. iioo). (Specimens 

determined by Burmeister.) 
1891 (Apidae) Scudder: 682 (5873). 
1907 (Apidae) Burmeister ; Handlirsch : 892. 
193 1 Apidae, Hiinefeld ; Salt : 144. 

Distribution. Eocene : Baltic amber. 

Remarks. This entry has been wrongly attributed to Burmeister. It was 
merely the latter's list that was used by Hiinefeld. 


Superfamily APOIDEA (except Apidae) incertae sedis* 
Apoidea gen. et sp. indet. i 

i88ia Apidae ; Malfatti : 98. 

1891 (Apidae) Malfatti ; Scudder : 682 (5874). 

1907 (Apidae) Malfatti ; Handlirsch : 892. 

Distribution. Oligocene : Chiavone, Italy. 
Remarks. Poorly preserved. 

Apoidea gen. et sp. indet. 2 

1881 Apidae ; Scudder : 290. 

1883 Apidae ; Scudder : 280. 

1891 (Apidae - several) Scudder : 682 (5875). 

1907 (Apidae - mehrere) Scudder ; Handlirsch : 892. 

Distribution. Oligocene : Florissant. 

Remarks. Scudder reported that several species occurred, 'most of them badly 
preserved ; the largest appears to be a Bombus'. 

Apoidea gen. et sp. indet. 3 

1907 (? Apidae) ; Handlirsch : 893. 

Material. ? Hofmuseum, Vienna (Bosniaski Collection). 

Distribution. Miocene : Gabbro, Italy. 

Remarks. Poorly preserved bee with short angulated antennae, habitus related 
to 'Apis melisuga' (see p. 248). 

Apoidea gen. et sp. indet. 4 

1895 (Apidae n.g. resembling Eucera) ; Scudder : 122. 
1907 Apidae n.g. ? Scudder ; Handlirsch : 890. 

Distribution. Miocene : Oeningen. 

Remarks. Some confusion exists over this entry. There are two difficulties. 
Firstly, according to Scudder (1895) and corroborated by Handlirsch (1907), this 
specimen may first have been considered a representative of Bombus grandaevus, 
but was later relegated to 'a new genus of Apidae, in which the neuration more 
closely resembles Eucera . Secondly Cockerell (1908b) investigated a specimen, 
from Scudder's collection, which he referred to the genus Calyptapis. The doubt 
that arises here is whether or not Cockerell was referring to the entry of Scudder in 
his reference to a Eucera during his discussion of Calyptapis. See Bombus floris- 
santensis (p. 212). 

* See also Appendix, p. 256. 


Apoidea gen. et sp. indet. 5 

192S Apidae Leach ; Pongracz : 156. 

Distribution. Miocene (Sarmatian) : Piski, Hungary. 

]\Iaterial. Magyar Allami Foldtani Intezet, Budapest. 

Remarks. According to Pongracz, the shape of the wings and the pointed 
abdomen indicate Osmia ; the large, contiguous eyes, however, point to Apis. 
Since only the male of Apis has contiguous eyes, whereas the pointed abdomen is 
not characteristic of the male, the specimen cannot be referred to Apis. On account 
of the general unreliability of the diagnoses of Pongracz it is impossible to say to 
which group of insects the fossil belongs. 


(excluding Apidae)^ 

'Formicinae' (Meunier) 

*I920 Andrena tertiavia Meunier : 730, pi. i, fig. 4. 

1931 Andrena tertiavia Meunier ; Salt : 145. 

1936 Andrena tertiaria Meunier ; Statz : 260. 

1936 Formicinen ; Statz : 260. 

HOLOTYPE. Heimatmuseum, Siegburg an der Lahn. 

Distribution. Oligocene : Rott. 

Remarks. Venation largely effaced. Statz (1936) writes : 'Nach der Bildung 
des Kopfes und der Fiihler, sowie der Thoraxsegmentierung und dem Bau der 
Beine gehort das Insekt gemass der Ansicht von Bischoff, Berlin, zu den Formicinen.' 
['According to Bischoff of Berlin, this insect belongs to the formicoids on the basis 
of the structure of the head and antennae, as well as the segmentation of the thorax 
and leg anatomy.'] 

'Formicinae' (Cockerell) [nee Hey den) 

1907 Anthophora effossa Cockerell : 228. 
1907 'An Ant' ; Cockerell : 228. 

Distribution. Oligocene : Rott. 

Remarks. An ant, to be found among a series of specimens from Rott, in the 
Mus. Comp. Zool. Harvard. 

Polistes kirbyanus Cockerell 1914 

1909c 'Three species of Osmia' ; Cockerell : 316 (partim). 

1909c A Wasp ; Cockerell : 316. 

♦1914 Polistes kirbyanus Cockerell : 638. 

1914 Osmia kirbyana Heer MS ; Cockerell : 638. 

^ See also Appendix, p. 257. 


HoLOTYPE. Eidgenossische Technische Hochschule, Zurich, No. PI. I, 681, with 

Distribution. Miocene : Oeningen. 

Remarks. The view is taken that among the 'three species of Osmia' referred to 
by Cockerell (1909c) as being at Zurich, one, the wasp, was called erroneously 'Osmia 
kirhyana . Not only was the generic identification wrong, but Heer appears to have 
given this specimen merely a manuscript name. 

'Genus' CELLIFORMA Brown 1934 

*I934 Celliforma Brown : 539. 
1935 Celliforma Brown : 527. 

Diagnosis. Fossil fillings of chambers purported to have been made by unknown 
digging or nest-building Hymenoptera. 

Genotype. Celliforma spirifer Brown 1934 by monotypy. 

Distribution. Cretaceous to Recent. 

Remarks. The name can hardly be maintained under the International Rules of 
Zoological Nomenclature, since it refers to objects made by animals, not to the 
animals themselves. It is used by Brown merely for the sake of convenience, in 
conformity with the practice adopted in palaeontology for fossil burrows, worm 
casts and footprints. 

Celliforma favosites Brown 1941 

*i94ia Celliforma favosites Brown : 54, pi. i, figs i, 2. 
1941 Celliforma favosites Brown ; Bequaert & Carpenter : 51. 
1941b Celliforma favosites Brown ; Brown : 105. 

HoLOTYPE. U.S. Geological Survey Collection. 

Distribution. Upper Cretaceous, Colorado and Montana Beds : Gunlock, 
south-west Utah, U.S.A. 

Remarks. Regarded by Brown as the comb of a wasp's nest resembling that of 
Polistes, but the possibility that it belongs to a social bee cannot be entirely dis- 
missed. Bequaert & Carpenter (1941) questioned the correctness of Brown's deter- 
mination, to which Brown (1941b) appears to have satisfactorily replied. 

Celliforma spirifer Brown 1934 

1934 Upper Eocene fossil molds ; Brown : 534, fig. 3. 
*I934 Celliforma spirifer Brown : 539. 

1935 Celliforma spirifer Brown ; Brown : 527. 

HoLOTYPE. U.S. Geological Survey Collection. 

Distribution. Eocene, Bridger Formation : Wyoming, U.S.A. 


Remarks. This cell resembles that of an Anthophora, with a rounded base and a 
spiral seal. Brown, however, emphasizes that many anthophorid bees construct 
such cells, hence little systematic significance can be attached to these features, 
except that they render probable the existence of Anthophoridae or similar bees. 

Celliforma germanica Brown 1935 

1907 Anthophora {Podalirius) ? sp. Schiitze : 25, pi. facing p. 56, figs 22, 23. 
♦1935 Celliforma germanica Brown : 527. 

Lectotype. Wiirttembergische Naturaliensammlung, Stuttgart (Schiitze's fig. 
22), here selected. 

Other specimens. Same collection, Schiitze's fig. 23, and a third specimen 
referred to on p. 26. 

Distribution. Oligocene, Bunter Breccia : Wilheim, Ries, Wurttemberg. 

Remarks. Schiitze recognized these as burrows of Hymenoptera, and thought 
they belonged to a species of Anthophora, but it is impossible to assign them to any 
particular genus. It should be noted that they differ from C. spirifer and C. nuda 
in the presence of a slight waist-like constriction. A spiral lid is not preserved. 

Celliforma nuda Brown 1935 

1915 Lithophaga nuda Dall : 129, pi. 24, fig. 4. 
1934 Lower Miocene fossil molds ; Brown : 534, fig. i. 
♦1935 Celliforma nuda Brown : 527. 

HoLOTYPE. U.S. Nat. Mus., No. 165187 or 165189 (Dall 1915 : pi. 24, fig. 4). 

Distribution. Lower Miocene, Silex beds of Tampa Limestone : Florida, U.S.A. 

Remarks. Originally described (Dall 1915) as moUuscan boreholes, but recog- 
nized by Brown (1935) as hymenopterous. Dall had both moUuscan shells of a 
Lithophaga nuda which he had described previously, and also burrows which he 
assigned to this species. Brown rightly identified as hymenopterous the burrow- 
filling figured by Dall ; hence it is the holotype of a species Celliforma nuda Brown 
(nee Dall). 

The cell resembles that of C. spirifer Brown in its structure, including the spiral lid. 

Celliforma bedfordi sp. no v. 

(PL I, figs 7, 8) 

Diagnosis. The innermost surface of the seal, that facing the main chamber, is 
distinctly spiral in structure, of at least five whorls, and is markedly concave. 
Holotype. British Museum (Natural History), In.31434. 
Type locality. Venus Bay, South Austraha. 

Distribution. Pleistocene or Sub-Recent, calcareous travertine and consolidated 
dune-rock, coasts of south and west Australia. 


Remarks. Specimens of these chambers were obtained, in 1933, by the British 
Museum (Natural History) from Mr R. Bedford of Kyancutta Museum, South 
Australia. They differ from those described by Lea (1925) in that they are tubular, 
with septate cross walls, some 6-7 cm long by 2 cm in diameter. According to the 
correspondence of Mr Bedford, 'the prevalent type is only the main chamber of a 
structure of which the missing portion consists of a considerable number of small 
chambers separated by concavo-convex septa'. The latter, however, are believed 
to be the various layers of a single seal to the main chamber. 

Celliforma septata sp. nov. 
(PL I, fig. 9 ; PI. 2, fig. i) 

Diagnosis. The innermost surface of the seal, facing the main chamber, is a 
plug which is not spiral. It appears to be just slightly convex. Seal composed of 
four septa. 

HoLOTYPE. British Museum (Natural History), In. 31435. 

Paratype. Same collection, In.34133. 

Type locality. Venus Bay, South Australia. 

Distribution. Pleistocene or Sub-Recent, calcareous travertine and consolidated 
dune-rock, south and west Australia. 

Remarks. These chambers appear to correspond to C. bedfordi n. sp. in size and 
shape, but differ in the form of the seal. 

Celliforma sp. indet. 
1932 'Tubular burrows in natural chalk' ; Buxton : 3. 

Described specimen. British Museum (Natural History), Dept. of Entomology. 

Distribution. ? Bronze Age, not later than King Solomon, Megiddo, Palestine. 

Remarks. Perhaps made by Osmia or Anthophora, according to Buxton (1932). 
No seal described. A second group of 'mud nests', submitted by Mr P. L. O. Guy, 
Oriental Institute, Chicago University, was also mentioned by Buxton. This 
appears to be a group of puparia reminiscent of the Australian forms described by 
Lea (1925), but somewhat smaller and made of different material. The latter 
were reasonably assumed to be the work of beetles still living in the vicinity. No 
such conclusion can be hazarded of the organisms responsible for the 'mud nests' 
of Megiddo. 

'Genus' URUGUAY RoseUi 1939 

Uruguay auroranormai Roselh 1939 

♦1939 Uruguay auroranormai Roselli : 73, figs 17-20, 22, 31(1-4). 
1941 Uruguay auroranormai Roselli ; Bequaert & Carpenter : 53. 
1941b Uruguay auroranormai Roselli ; Brown : no. 


Distribution. Cretaceous or later : Uruguay, 

Remarks. Unfortunately the original work in which Roselli described the 
hymenopterous cells has not been seen, but it is fairly obvious that Bequaert & 
Carpenter's comparison of Brown's Cretaceous fossil, Celliforma favosites (p. 203), 
with U. auroranormai is far-fetched. It would appear from the descriptions of 
Bequaert & Carpenter (1941) and Brown (1941b) that the cells are not dissimilar to 
those described in the present work from south and west Australia and may well be 
the work of bees. Moreover, as Brown has pointed out, the fact that they occur in 
Cretaceous strata does not imply a Cretaceous origin for them. 


This section contains a detailed study of the fossil Apidae. The material is 
arranged according to tribes, and within them stratigraphically. 

Family APIDAE Latreille 1802 

Diagnosis. Social bees, except possibly the tribe Euglossini (which has not so 
far been found in the fossil state) ; second flagellar segment much shorter than scape ; 
basitibial plates absent ; scopa of female, when present, forming a corbicula on each 
posterior tibia, except in queens of Meliponini ; inner apical margins of posterior 
tibiae of non-parasitic females, except in queens of Meliponini and workers of 
Lestrimelitta, provided with a tibial pollen rake ; pygidial plate absent ; wings hairy 
throughout, except possibly in a few fossil forms ; usually (and in all North American 
forms) the distance between the costal ends of the two recurrent veins is nearly twice 
as great as the length of the second recurrent vein, or even more, and is longer than 
the first recurrent. 


Many fossil insects carry characters of such a type that their exact systematic 
position is difficult to define. Often new families, subfamilies and tribes are created 
for them but not infrequently this procedure defeats its own end by establishing 
units of separation where none exist. Chalcobombus embraces such a group and it 
seems to us inadvisable to erect a new tribe for it. This genus could be regarded as 
having affinities with the Bombini, or else it may have been the progenitor of 
Sophrobombus which we consider to be an ancestral Meliponine, and thirdly, it is not 
without affinity with the genus Electrapis which links it with the Apini. For these 
reasons we have not assigned a tribal name to the genus. 

Genus CHALCOBOMBUS Cockerell 1908 

* 1 908b Chalcobombus Cockerell : 327. 
igogd Chalcobombus Cockerell ; Cockerell : 12. 

Genotype. Chalcobombus humilis Cockerell 1908, by subsequent designation of 
Cockerell (i909d : 11). 


Description. Apidae of compact build, about 8 mm long ; metallic tints 
sometimes in evidence ; head broad, vertex usually with coarse, erect bristles, 
sparingly plumose, facial quadrangle variable ; eyes not usually hairy, and ocelli 
large, close together, in a slight curve ; scape comparatively long, flagellum variable 
in size ; mouth parts elongate, 'tongue' hairy but slender with an apical circular 
flabellum ; mandible with at least two marginal notches ; abdomen subglobose ; 
scutellum prominent and hairy ; legs hairy, hind ones considerably plumose, 
pul villus large, claws with short inner tooth ; antenna cleaner with exceptionally 
long hairs, lining the notch or 'auschnitt', its clasp unusually large, fringed with 
very fine hairs along its inner surface, lacking a lobe, variable in size ; along the 
distal margin of the hind tibia a crude rake of strong stiff setae, and flanking its 
anterior margin an inner tibial spur with one edge of stiff hairs, closely set. This 
spur is fairly closely pressed against the legs. 

Forewing. Stigma distinct, slightly variable in size, triangular ; marginal cell 
deepish, not particularly long ; three subequal submarginal cells, the first sometimes 
the largest, but all three exhibiting variations in shape that are of specific value ; 
second submarginal cell receiving first recurrent nervure beyond or near the middle ; 
third submarginal cell receiving second recurrent nervure a short distance from its 
end ; junction of cubito-anal cross vein and basal nerve coincident. 

Hindw^ing. Cross vein cu-a somewhat oblique, or the upper half vertical ; 
hamuli variable in number (8- 11). 

Remarks. The genus was only mentioned in 1908 in a list of Chalcohomhus 
species and in a genealogical tree. It was fully described in 1909. The above 
description is based entirely on Cockerell's published papers. 

Chalcobombus hirsutus Cockerell 1908 

♦1908b Chalcohomhus hirsutus Cockerell : 326. 
igogd Chalcohomhus hirsutus Cockerell ; Cockerell : 12, text-fig. 8. 

HoLOTYPE. Geological Museum, University of Koenigsberg. 

Distribution. Eocene : Baltic amber. 

Diagnosis. Head and thorax with long whitish hairs. 

Description. A compact black bee, 8 mm long ; head and thorax, particularly 
the scutellum, with long, white or yellowish hairs apparently intermixed with brown 
ones, and all sparingly plumose ; face, particularly below, broad ; scutellum con- 
spicuously projecting ; wings reddish-fuliginous ; distal corner of second sub- 
marginal cell produced, third submarginal cell longer than second, narrowed more 
than half above. 

Basitarsus as in C. humilis ; legs with much coarse fuscous hair ; flagellum is 
given as about 0-255 "ini long and 0-170 mm thick, last joint 0-323 mm long. 

Remarks. This species was first listed with characters in 1908, but not fully 
described until 1909. 


Chalcobombus hutnilis Cockerell 1908 

* 1908b Chalcobombus humilis Cockerell : 326. 
igogd Chalcobombus humilis Cockerell ; Cockerell : 12, text-figs 6, 7. 
1 93 1 Chalcobombus humilis Cockerell ; Salt : 145. 

HoLOTYPE AND Paratype. Geological Museum, University of Koenigsberg. 

Distribution. Eocene : Baltic amber. 

Diagnosis. Head and thorax above with short, black hair. 

Description. Black, about 8-5 mm long ; head and thorax above with short, 
black hair ; vertex and front dull, with sparse, short, coarse, erect, black bristles, 
sparingly plumose ; second 'antennal joint' barrel-shaped ; wings dark fuliginous ; 
stigma triangular and distinct, marginal cell ending rather obtusely away from costa 
and possessing a small appendiculation ; second submarginal cell greatly narrowed 
above, narrower on marginal than is the third ; third submarginal cell narrowed at 
least half above. 

Basitarsus squarish ; flagellum a little variable in size and possibly in thickness - 
Cockerell quotes lengths of 1-955 mm and 2-125 ^^^^, and thickness 0-187 ni^i- 

Remarks. This species, selected by Cockerell (i909d) as the genotype, was 
first listed with characters in 1908 but not fully described until 1909. 

Chalcobombus martialis Cockerell 1908 

*i9o8b Chalcobombus martialis Cockerell : 326. 
i909f Chalcobombus martialis Cockerell ; Cockerell : 22. 
193 1 Chalcobombus martialis CockeTell ; Salt : 145. 

HoLOTYPE. Geological Museum, University of Koenigsberg. 

Distribution. Eocene : Baltic amber. 

Diagnosis, Head, mesothorax and scutellum dull black, prothorax and tubercles 
coppery red. 

Description. About 8 mm long ; mesothorax, scutellum and vertex dull black, 
but prothorax and tubercles coppery red ; face and abdomen black but legs metallic 
copper-red or crimson, moreover, the hind margins of the segments very narrowly 
but conspicuously silvery white, the dark part next to the margins stained with red, 
especially on the first segment ; scape shortish ; hind basitarsus twice as long as 
broad ; third submarginal cell is broad above ; first recurrent nervure joins second 
submarginal cell near middle ; wings pale reddish brown ; stigma large but not of 
great width ; second recurrent nervure with strong double curve. 

Thick-set bee ; pubescence black or dark fuscous, not abundant ; the antennal 
joints appear to be known with some precision, as follows : third longer than second, 
fourth very short, about twice as broad as long, fifth conspicuously longer than 

Remarks. This species was listed with characters in 1908, but not fully described 
until 1909. 


Tribe BOMBINI Latreille 1802 

Diagnosis. Moderate-sized to large hairy bees ; clypeus slightly protuberant, 
its lower lateral portions being slightly bent backward ; malar space long ; labrum 
about twice as broad as long ; proboscis usually not reaching beyond middle coxae ; 
scutellum rounded ; posterior tibiae of male neither swollen nor fossate ; two 
posterior tibial spurs always present ; pterostigma often small, not tapering beyond 
base of vein r ; apex of marginal cell separated from wing tip by a distance equal 
to at least half of cell ; jugal lobe of hindwing becoming reduced or absent. 

Remarks. No primitive forms are known of this tribe, the earliest appearing in 
the OUgocene (Aquitanian), and these are poorly preserved. The first convincing 
bumble bees are of Oligocene age, from Florissant. [For Probombus see Appendix, 
p. 256.] 

Genus BOMBUS Latreille 1802 

*i8o2a Bombus Latreille : 437. 

1906 Calyptapis Cockerell : 42 (Genotype Calyptapis florissantensis Cockerell 1906 by 

1908b Calyptapis Cockerell ; Cockerell : 324. 

Genotype. Apis terrestris Linnaeus 1758 [ICZN Opinion 220, 1954]. 

Diagnosis. Moderate-sized to large hairy bees ; clypeus slightly protuberant ; 
female with well-developed corbicula on each hind tibia, the latter also possessing 
two strong spurs but no developed comb. 

Basitarsus rectangular, strongly developed and inner surface covered with strong 
coarse bristles ; depstum or auricle present. Posterior tibia of male flattened but 
without corbicula. 

Bombus crassipes Novak 1877 

♦1877 Bombus crassipes Novak : 92, pi. 3, fig. 4. 

1891 Bombus crassipes Novak ; Scudder : 690 (5951). 

1907 Bombus crassipes Novak ; Handlirsch : 891. 
1928 Bombus crassipes Novak ; Wheeler : 97. 
193 1 Bombus crassipes Novak ; Cockerell : 309. 
1936 Bombus crassipes Novak ; Statz : 260. 

HoLOTYPE. Whereabouts unknown. 

Distribution. Oligocene (Aquitanian) : Krottensee, Bohemia. 

Description. Length of body 10 mm, width 6 mm ; dark brown in colour, 
densely hairy ; hind legs strongly developed, very hairy ; auricle possibly not so 
well developed ; femurs of hind legs thick ; tibiae flat, widening posteriorly and 
truncated ; basitarsus large, flat and truncated at both ends. 

Remarks. One specimen only, not well preserved, neither head nor wings being 
discernible. Fortunately Novak's description is helpful. It suggests that the speci- 
men was a genuine Bombus. The figure, however, is very bad and possibly incorrect 


since three hind legs are displayed. What may be the right hind leg is in the correct 
position, and except for the omission of the auricle and the tibial spurs is remarkably 
reminiscent of Bomhus. Of the left hind leg, the basitarsus only is satisfactorily 
preserved. This, too, is Bomhus-Vike. The third 'leg' is depicted in an anterior 
position where in fact the wing might be expected ; perhaps only a mass of chitin is 
intended, which has been given the semblance of a basitarsus by the artist. The 
hind leg is very hairy and the corbicula is clearly indicated. 

Bombus abavus Heer 1867 

(PI. 2, fig. 2) 

*i867 Bomhus abavus Heer : 5, pi. 3, figs 9, 10. 

1891 Bomhus abavus Yieer ; Scudder : 690 (5948). 

1907 Bombus abavus Heer ; Handlirsch : 892. 

1909c Xylocopa abavus (Heer) ; Cockerell : 314. 

1928 Bombus abavus Heer ; Wheeler : 97. 

1928 Xylocopa abavus (Heer) ; Wheeler : 97. 

1 93 1 Bombus abavus Heer ; Cockerell : 309. 

1 93 1 Xylocopa Cockerell : 309. 

1938 Bombus abavus Heer ; Armbruster : 87. 

HoLOTYPE. Said to be in Eidgenossische Technische Hochschule, Zurich, but no 
trace can be found in that collection, and it must be presumed lost. 

Material. Two specimens in the Eidgenossische Technische Hochschule, 
Zurich (Nos. PL I, 808, examined by Cockerell, and SN.201) ; one specimen in the 
Badische Landesammlung fiir Naturkunde, Karlsruhe (No. Inv.1268) (Oe.90)). 

Distribution. Miocene, Oeningen. 

Description and Remarks. Cockerell and Armbruster, who did not examine 
the type, were inclined to place this species in Xylocopa, possibly because of Heer's 
remark that the basitarsus is hairy. The venation, however, which Cockerell 
wrongly maintained could not be made out, is somewhat like that of Bomhus in 
specimen No. PL I, 808. Moreover, the 'globular' abdomen of specimen SN.201 is 
not a feature of Xylocopa. In the circumstances it is advisable to retain this form 
in the genus in which it was originally placed, but its systematic position remains 
uncertain. The three existing specimens may be described as follows. 

Specimen PL I, 808. Only thorax and legs are preserved with parts of the 
wings, especially the greater part of one forewing. The basal vein is fairly long, and 
its posterior segment is much longer than its anterior segment ; cross vein cu-a is 
slightly distad of junction of basal nerve with Cu ; second recurrent nerve is angular 
shortly before it meets the base vein of the second submarginal cell. The last 
recurrent nerve is incompletely preserved, and the junction lies in the proximal 
half of this vein. 

The wing differs from that of Xylocopa in that the anterior segment of the basal 
vein, although short, is not so short as is usual for this genus, secondly in the com- 
paratively slightly greater length of the basal vein, and thirdly in the fact that the 


second recurrent nerve meets the base vein of the second submarginal cell well away 
from its apex. In these characters it resembles Bombus. 

Forewing length at least 6-5 mm. The legs/ particularly the hind ones, are stout 
and hairy but not so much as to warrant the exclusion of the species from Bombus. 
The broad thorax reveals substructures of the notum, which was not without hairs. 
Length of thorax 4 mm, width 5-5 mm. 

Specimen SN.201. Head not preserved. The fossil consists of thorax and 
abdomen with parts of legs, and the merest trace of a wing. The thorax and abdomen 
are hairy and there is a strong suggestion of hairs on the legs. Since the type 
figured by Heer indicated an insect which had part of the head and several legs 
present, it is obvious this is not Heer's type. Length of thorax 4 mm, width 5 mm ; 
length of abdomen 6 mm. 

Specimen Inv.1268 (Oe.90). This specimen was identified on the label as 
Xylocopa senilis. Like the other known fossils referred to this species (p. 191) it is 
poorly preserved, lacking head, part of thorax, and all legs except one hind femur 
and tibia. The abdomen, however, is complete, and the hairs and segmentation 
very distinct. Moreover, wings are present, that on the right showing some im- 
portant features of the venation. Dimensions : length of abdomen 10-5 mm, 
width 7 mm ; length of forewing 10 mm ; length of hind tibia 3-5 mm. It is thus 
clear that the specimen is larger than the other known specimens of B. abavus, but 
since bumble bees vary much in size, this is no reason for excluding it from the present 

Diagnostic characters are seen in the shape of the marginal cell, particularly in 
the concavity of its base, which in Xylocopa is convex and very acute proximally, 
in the position of the insertion of the first submarginal cross vein (abscissa Rs) on 
the posterior margin of the marginal cell which is much more distal in Xylocopa, and 
also in the curve, or rather indentation, of the cubitus just prior to its distal splitting 
into Cui and Cua, a character much less prominent or lacking in Xylocopa. These 
are not conclusive arguments for placing the specimen in the genus Bombus rather 
than Xylocopa, but since in addition the long tibia has strong hairs arranged in a 
manner suggesting a corbicula, the evidence becomes overwhelming. That femur 
and tibia are preserved, rather than tibia and basitarsus, is proved by the angle 
between them. The basitarsus, indeed, is present also but only very incompletely 
preserved ; superficially it resembles the distitarsi. Its erroneous interpretation as 
distitarsi no doubt contributed to the earlier identification of the specimen with 

The colour of the chitin and hairs is dark brown. Hairs are very conspicuous on 
the apex of the abdomen but become much sparser on the base of the abdomen and 
the thorax. Markings on the thorax indicate the presence of morphologically 
distinct regions, but these are too poorly preserved to be of use for generic identifi- 

* Cockerell (193 1) maintained, 'The legs show a scopa, and the hind tibia is very broad, with a gently 
curved longitudinal ridge visible on both sides, and, therefore, certainly natural. This ridge is normal 
for Xylocopa, to which Bombus abavus Heer must apparently be referred'. This ridge, however, could 
quite well, in part, be the outline of the corbicula of a Bombus, and in any case would almost certainly 
have been masked by the scopa in a fossil Xylocopa. 


Bombus florissantensis (Cockerell 1906) 

1895 Apidae n.g. ; Scudder : 122. 
*i9o6 Calyptapis florissantensis Cockerell : 42. 
1907 Calyptapis florissantensis Cockerell ; Handlirsch : 890. 
1908b Calyptapis florissantensis Cockerell ; Cockerell : 324, 326. 
1909b Calyptapis florissantensis Cockerell ; Cockerell : 80. 
193 1 Calyptapis Cockerell ; Cockerell : 311. 

HoLOTYPE, Mus. Comp. Zool. Harvard, No. 2008 (= Scudder Coll. No. 4933). 

Material. Specimen studied by Cockerell (1908), from the Oligocene shales of 

Distribution. Oligocene : Florissant. 

Diagnosis. Length of body 15 mm ; length of anterior wing 8-5 mm ; head and 
thorax black but abdomen rather pale reddish, the junctions of the segments dis- 
tinguished by moderately broad, light bands ; although legs are hairy, as in Bombus, 
and the hind legs exhibit a corbicula, the abdomen is not noticeably hairy, and the 
inner tooth of the bifid claws is much smaller and shorter. 

Wings hyaline, with pale veins, cubito-anal cross vein (cu-a) of forewing vertical 
except where it bends away immediately prior to the junction with basal nerve. It 
differs from Recent Bombus 'in the form of the third submarginal cell and in the 
somewhat less specialised second submarginal' (Cockerell 1908b). 

Stout-bodied, like Recent Bombus ; hind basitarsus flattened and quadrate, with 
hairy margins. It is stated to be 'broadly emarginate apically'. 

Remarks. The above description is based on Cockerell's of 1908. He placed the 
specimen in the genus Calyptapis because of the peculiarities of the second and third 
submarginal cells. Apart from the vagueness of his statement, however, it is 
precisely at this part of the wing that variation might be expected in an Oligocene 
species. The erection of a new genus indeed appears to confuse the issue, for it 
seems to us that we are here dealing with a perfectly legitimate Bombus able to bear 
the most detailed comparison with Recent forms. 

Formerly Cockerell regarded it as near Melissodes (Anthophoridae), but since 1908 
as one of the Bombini. He apparently based his first description on a poorly pre- 
served specimen already referred to Eucera by Scudder (1895), who said it was 'an 
insect which may possibly be Bombus grandaevus Heer, but which belongs to a new 
genus of Apidae, in which the neuration more closely resembles that of Eucera'. 
Later, a better specimen was discovered which clearly indicated the relationship 
with Bombus. 

Bombus proavus Cockerell 1931 

♦1931 Bombus proavus Cockerell : 309, pi. i, fig. 8. 
1936 Bombus abavus Cockerell ; Statz : 295. 

Holotype. Mus. Comp. Zool. Harvard, No. 2940a, b. 

Distribution, Miocene, Latah Formation ; near Spokane, Washington, U.S.A. 


Diagnosis. A robust insect, with long black hair, at least on the face ; forewing 
15 mm long ; basal vein slightly proximal of junction of vein cu-a ; first submarginal 
cross vein (abscissa Rs) not angulate ; first marginal cell on first media cell slightly 
longer than second submarginal on same cell ; second submarginal cell on the 
marginal cell very little longer than the third on same cell ; lower side of third sub- 
marginal conspicuously bent at insertion of second recurrent ; wing hyaline, slightly 
brownish, the upper part of the marginal cell moderately infuscated. 

Remarks. Insect known from Cockerell's description and figure. Its body is 
so crushed that most features are obliterated, but the forewing is in an excellent 
state of preservation. As with B. florissantensis (Cockerell) the third submarginal 
cell in this specimen differs from Recent Bombus. 

According to Cockerell this was the first true fossil Bombus to be described, and 
all other species belong to other genera or are of doubtful reference to Bombus. He 
also maintains that it is 'nearest to European B. lapidarius' . Statz (1936) when 
referring to this specimen erroneously named it B. abavus. 

Tribe MELIPONINI Handlirsch 1924 

Description. Moderate-sized to minute bees, in perennial colonies ; Recent 
forms with eyes bare, sparsely hairy in some queens ; labrum much broader than 
long ; epistomal suture arched upward, as in Apis, close to the antennal sockets, 
and the clypeus not protuberant. 

Posterior tibial spurs reduced or absent ; tibial comb always present in Recent 
forms (Lestrimelitta excepted) and distinct ; in fossil forms if a comb is not present 
there is always an inner but modified tibial spur. Basitarsus very rarely as broad 
as tibia ; auricle or true depstum apparently always absent, while the inner surface 
of the basitarsus may not have the whole area completely covered with strong 
bristles ; claws of Recent females simple. 

Ninth sternum of male virtually absent, being represented, if at all, by a narrow, 
longitudinal, weakly sclerotic band beneath the genitalia ; eighth sternum rather 
narrow, having the form characteristic of the ninth in most bees ; sting reduced. 

Wings extending beyond the body ; marginal cell often open, and forewing with 
veins always weak (or weakening) distally ; pterostigma of moderate to large size, 
extending well beyond base of vein r ; jugal lobe of hindwing present. 

Remarks. The genus Sophrobombus is represented by fossil forms, but not 
Melipona. The tribe ranges from Eocene to Recent and is today widely distributed 
in the tropics. Many are found in East African copal. 

Genus SOPHROBOMBUS Cockerell 1908 

•1908b Sophrobombus Cockerell : 327. 
igogf Sophrobombus Cockerell ; Cockerell : 21. 

Genotype. Sophrobombiis fatalis Cockerell 1908, by monotypy. 


Diagnosis, Small bees resembling Chalcobombus ; mandibles with at least a 
convex cutting edge ; ocelli large and arranged in a curve. 

Basitarsus flattened, quadrangular, short, having on the inner surface a small, 
bristleless area around the posterior apex ; a ridge of short hairs exists on the inner 
surface of the hind tibia ; inner hind tibial spur present, fringed with hairs along 
one edge, and acting as a 'stop' to the pollen rake. 

Forewing with second submarginal cross vein absent ; basal vein going basad 
some distance from vein cu-a ; pterostigma well developed ; marginal cell possibly 
slightly open at apex. 

Remarks. Cockerell considered this genus to be a near descendant of Chalco- 
bombus. It differs from it, however, in a number of points, particularly in the reduc- 
tion of the wing venation. In this respect it is much nearer Trigona which it closely 
resembles in habitus, as well as in the hairless patch on the inner surface of the 
basitarsus and the ridge on the inner surface of the hind tibia. There seems little 
doubt therefore that it could have been an ancestor of the Meliponini. The presence 
of the inner tibial spur, rather than hindering this view, fosters it. 

The genus was first mentioned by Cockerell (1908) in a list of Baltic amber species 
and in a genealogical tree. It was fully described in the following year, 

Sophrobombus fatalis Cockerell 1908 

(PI. 2, figs 3, 4) 

* 1908b Sophrobombus fatalis Cockerell : 326. 
igogf Sophrobombus fatalis Cockerell ; Cockerell : 21. 
1931 Sophrobombus fatalis Cockerell ; Salt : 146. 

HoLOTYPE, Geological Museum, University of Koenigsberg, 

Material, Scheele Collection, No, 1225, in Geologisches Staatsinstitut, Hamburg, 

Distribution. Eocene : Baltic amber. 

Diagnosis. Length 8-8-5 i^^i- Short, thick-set insect, metallic dark coppery- 
red, almost without pubescence. Vertex with very short fuscous plumose hair. 
Mandibles broadened apically, with a convex cutting edge ; clypeus weakly punctate ; 
scape comparatively long and curved. Thorax above with erect, strongly plumose 
fuscous hair, twice as long on scutellum as on mesothorax ; scutellum projecting. 
Hind tibia long and large ; hind basitarsus quadrangular, less than twice as long as 
broad, the outer angles acute. Claws with an inner tooth a little beyond the middle. 

Description. Cockerell gave the following description of the specimen : 'Man- 
dibles (apparently without visible teeth) with long and short bristles on the lower 
margin ; blades of maxillae long and slender ; antennae very much closer together 
than either is to the eye, second joint barrel-shaped, but its basal third narrower 
and cylindrical, third joint much longer than fourth but not quite so long as second, 
fourth broader than long ; legs moderately clothed with fuscous hair ; pulvillus 
large ; 2-4 tarsal joints, small, cordiform ; wings pale fuscous, veins and stigma 
dark brown ; hind wing with vein cu-a a little oblique, with the lower endmost 
apicad.' We have examined specimen no. 1225 and find that it agrees closely with 


the above description, except that the second flagellar joint, i.e. third antennal 
joint, is somewhat longer than the first. This does not, however, justify creating a 
new species. The detailed description of this new specimen is as follows. 

A metallic, coppery-red insect, not hairy. Overall length, from head to wing tip, 
8 '5 mm ; the actual body, however, is shorter, with an overall length from head to 
the tip of the abdomen of only 6 mm. The extent of wing overlap is thus consider- 
able but was possibly somewhat less in the living creature since the whole abdomen 
is compact, which suggests considerable contraction. Other approximate measure- 
ments are : width of thorax 275 mm, length 2-5 mm ; width of head 2 mm, length 
1-25 mm ; length of tibia 2-5 mm. 

Head. The rather large head is not preserved in its entirety since a hollow occurs 
in the amber near the mouth parts, and some 'schimmel' also has developed here. 
The upper part, however, is distinct. The compound eyes appear to be naked ; 
the vertex is covered with fuscous hair, possibly plumose ; ocelli are large and 
prominent, set in a broad shallow triangle ; malar space quite broad but short, if 
the limits of observations are the actual limits of this region ; one antenna quite 
distinct, inserted on the lower half of the face ; scape well formed, curved ; basal 
segment of flagellum, i.e. the pedicel, barrel-shaped, second slightly larger than first, 
but tapering basally, third the shortest, broader than long but not much shorter 
than the fourth segment, the terminal segment the largest, at least twice the size of 
the remaining segments which are equal in shape and size. Eleven flagellar segments 
are present. 

Thorax above indistinct owing to cracked amber and some 'schimmel'. It was 
apparently well arched and covered with hairs which extended somewhat down the 
sides. The projected scutellum can be observed. 

Abdomen. Ovoid in shape. Although the tip of the abdomen can be seen very 
distinctly, there is no trace whatever of a sting. 

Legs. Only the hind legs have been sufficiently preserved for study. They are 
remarkable for the great length of the tibia with a shallow scoop-like extension over- 
lapping the base of the basitarsus on the outside, and the comparatively small, 
quadrangular basitarsus, not much longer than broad, with the posterior distal 
angle strikingly acute. In detail, the hind leg has the following characters. 

Tibia. Corbicula present with the usual scopa, some hairs of which may be 
slightly plumose ; the corbicula itself relatively free from hairs ; the inner surface 
of the tibia exhibits, posteriorly, a longitudinally arranged plateau-Uke region of 
short close hairs, occupying a third of its area and extending slightly to the femur, 
and bounded (though seen only under certain lighting conditions) by a dark rim 
which is quite broad towards the base of the femur but narrow on the tibia except 
for a broader zone at the pollen rake. Two appendages are to be found on the 
tibia, a strong pollen rake and an inner tibial spur, the latter having become modified 
to carry a blade of fine hairs, as though developing into a comb. That it is not the 
latter is clear from its position on the inner surface of the tibia where it forms a 
'stop' to the pollen rake. It also differs from the typical spur by being more closely 
pressed against the leg and more intimately associated with the pollen rake. 


Basitarsus. The inner surface exhibits a well-developed pollen brush, consisting 
of seven rows of strong bristles, with possibly 18-20 bristles to the middle rows, 
though the latter number is difficult to ascertain with complete accuracy owing to 
pollen clogging the brush. Apical and basal rows have at least four bristles less in 
consequence of the development of small, bristleless areas around the posterior 

Pidvillus large ; claws bulbous at base and strong, with very small tooth beyond 
half-way ; remaining tarsal joints cordiform. 

Wings. Fuscous, hairy, and rather more elongate than is normal for the Baltic 
amber bees. A study of the venation of the fore wing reveals the absence of the 
second cross vein of the submarginal cells ; the basal vein becoming basad a short 
distance away from vein cu-a ;' the latter vein a little oblique, the posterior end 
most apicad ; stigma well developed ; marginal cell pointed ; second submarginal 
cell shorter than slightly open marginal ; second recurrent vein weak and sloping 
strongly basad. Since the fore wings are folded along the back of the insect, the 
hindwings are not visible. 

Remarks. The absence of reference by Cockerell to denticles on the mandible 
cannot be taken too seriously since such denticles, unless very prominently displayed 
on an open mandible, are not easy to see. Cockerell's specimen was listed with 
characters in 1908, but not fully described until 1909, 

Genus TRIGONA Jurine 1807 

♦1807 Trigona Jurine : 246. 
1896 Meliponorytes Tosi : 352 (type species Meliponorytes succini Tosi). 

Genotype. Apis amalthea Olivier 1789, by subsequent designation of Latreille 
(1810 : 439). 

Diagnosis. Stigma well developed, rounded below ; wings relatively long, 
extending beyond apex of abdomen ; hamuli about five or six, rarely more than nine 
or fewer than four ; body length from 2 to 8 mm, rarely exceeding 11 mm. 

Remarks. The genus Meliponorytes was erected by Tosi (1896) to embrace forms 
ancestral to Recent Meliponini. All the fossils so far discovered and which have 
recognizable descriptions, however, are believed to be referable to Trigona. But 
the creation of the new genus Meliponorytes for what is apparently a genuine Trigona 
must have been a deliberate procedure, since Tosi makes mention of the likeness of 
his specimens to Trigona and also to the subgenus Tetragona. Meliponorytes was 
thus undoubtedly erected on account of those slight uncertainties attending fossil 
forms, although the action here seems scarcely justifiable. Tosi's descriptions and 
illustrations have been very carefully studied and there seems little doubt that his 
M. succini (see p. 221) is definitely a Trigona of the subgenus Tetragona, whilst the 
subgenus cannot be stated for his M. sicula (p. 224). Kerr & Maule (1964) thought 

' Not, as stated by Cockerell for his specimen, a considerable amount, though it is the most we have 
noticed among Baltic amber bees. 



that M. succini belonged to Meliponula, but in a different subgenus from M. 

It will be recalled that all Trigona species lack hind tibial spurs, yet all, except 
Lestrimelitta, possess the hind tibial comb. The variations of the basitibial surfaces 
are of subgeneric value. [For subgenus Nogueirapis see Appendix, p. 257.] 

Subgenus TRIGONA Jurine 1807 

♦1807 Trigona Jurine : 245. 

Genotype. Apis amalthea Olivier 1789. 

Diagnosis. (From Schwarz 1948.) Mandible toothed usually from end to end 
along its apex, or with three teeth on at least the outer two-thirds of the apex and 
an angulation at the inner extremity. Hind tibiae a little longer than the combined 
length of their femora and trochanters. The fringe along the posterior lateral 
contour of the hind tibiae with plumose hairs in addition to the usually sparser 
simple hairs. The base of the inner face of the hind basitarsi with a differentiated, 
more or less oval, area of appressed hairs that contrast with the brush-like hairs on 
the apical half to two-thirds of this inner face. 

Remarks. At least one fossil form is so far known, from the Pleistocene. 

Trigona (Trigona) erythra Schletterer 1891 

*i89i Trigona erythra Schletterer : 2. 

1895 Melipona togoensis Stadelmann : 620. 

1948 Trigona erythra var. togoensis Stadelmann ; Schwarz : 51, 61, 68, 132. 

1961 Axestotrigona erythra (Schletterer) ; Moure : 239. 

1961 Axestotrigona togoensis (Stadelmann) ; Moure : 239. 

Fossil specimen studied. British Museum (Natural History), In.17664. 

Distribution. Pleistocene : East African copal ; and Recent. 

Description. Body length 5-75 mm ; general colour brownish-black to black. 
Head broader than thorax, hairs upright and relatively strong on vertex, ocelli 
prominent, in a gently curved line, hind ocelli more distant from each other than 
from compound eyes ; from the front ocellus there runs a prominent frontal line ; 
face, particularly the clypeus, appears to be covered with a close white pubescence ; 
length of scape little more than half the length of the flagellum, and all flagellar 
segments, except the last, not longer than broad, the last being longer than broad, 
and pointed. 

Thorax markedly triangular, black, its surface matt, probably due to a short 
pubescence and to very fine rugosity ; longitudinal line through scutum (median 
mesoscutal line) masked by a crack in the amber, but the polished and shining furrow 
between scutum and scutellum can just be recognized ; scutellum clearly projecting, 
with strong chitinized hairs on its ridge. 

Abdomen dark in colour with only few traces of brown, anterior segments smooth 
and shiny, hind segments covered with short pubescence ; legs on the whole black 


but not sufficiently well preserved at extremities to judge whether the latter are 
brown (most have been cut off short at the edge of the copal) ; corbicula well formed, 
wide, neatly hollowed, inner surface difficult to see ; one basitarsus detached, the 
other wanting ; claws not notched. 

Marginal cell of forewing a little open at apex ; one rather weak submarginal 
cross vein ; first recurrent distinctly angular before junction with vein M ; seven 

Remarks. No diagnosis is given because the taxonomic position of the Recent 
species is still sub judice, and we are not sufficiently acquainted with previous investi- 
gations concerning this group to be able to make a useful contribution. 

The fossil agrees with Recent specimens of Trigona togoensis Stadelmann in the 
British Museum (Natural History) and our description of the fossil agrees in all 
essential respects with the original description of T. togoensis by Stadelmann. 

The two Recent forms T. togoensis and T. erythra differ in the colour of the 
abdomen, which is rusty red in the latter. T. togoensis is also slightly smaller. 
Schwarz, however, regards these differences at best as of varietal value, and we have 
followed him in regarding them as races of a single species. In the colour of the 
abdomen, the fossil from East Africa agrees with the West African form T. togoensis. 

Subgenus Hypotrigona Cockerell 1934 

*I934 Hypotrigona Cockerell : 54. 
1961 Liotrigona Moure : 223 (type species Trigona bottegoi Magretti). 
1961 Hypotrigona Moure : 220. 

Genotype. Trigona gribodoi Magretti 1884, by original designation. 

Diagnosis. Minute bees, with dull tegument but without true yellow marks. 
Clypeus flat, a little wider than twice its length and one-third as long as clypeocular 
distance. Lateral portion of epistomal suture straight. Mandible bidentate. 
Scape short. Flagellar segments very short. Bifurcation between M and Cu after 
cu-an. Submarginal angle about 90°. Hind basitarsus about twice as long as 
broad. [See also Appendix, p. 256.] 

Trigona (Hypotrigona) gribodoi Magretti i^ 
(PI. 2, figs 5-7) 

•1884 Trigona gribodoi Magretti : 630. 

1894 Trigona braunsi Kohl : 280. 

1895 Trigona bottegoi Magretti : 156. 

1934 Trigona {Hypotrigona) gribodoi Magretti ; Cockerell : 54. 

1934 Trigona [Hypotrigona) bottegoi Magretti ; Cockerell : 55. 

1961 Hypotrigona gribodoi (Magretti) ; Moure : 220. 

1 96 1 Liotrigona bottegoi (Magretti) ; Moure : 223. 

1964 Trigona [Hypotrigona) gribodoi Magretti ; Kerr & Maule : 7. 

1964 Trigona [Liotrigona) bottegoi (Magretti) ; Kerr & Maule : 7. 


Specimens studied. British Museum (Natural History), In.38982- 38993, 
In.17686, In.18209, In.18214, In.18220, In.18226, 58511, 58622. 

Distribution. Recent (common in Africa) ; Late Pleistocene : East African 

Diagnosis. Length of body not exceeding 4 mm ; head and thorax black, 
abdomen and eyes dark brown. There is no apparent natural testaceous coloration 
of the scape, mandibles or anterior legs. The body is only sparsely haired except, 
chiefly, for the ridge of the scutellum and the basitarsi ; hind tibia triangular, neither 
clavate nor club-shaped, the inner face showing an elevated plateau-like area with a 
posterior rim. 

Description. Face round, black ; antennae of only moderate length, flagellum 
broad and shortish ; ocelli prominent. Although the whole face appears uniformly 
black, the copal is believed to mask its short white pubescence, except in In. 38993. 
The same is true of the hairs on the black thorax, but the stronger, more heavily 
chitinized hairs of the scutellum can be distinguished. Hind tibia not copiously 
haired, and because the hairs are fuscous they are sometimes difficult to see. All 
the tarsi appear to be more strongly haired than the tibiae. Wings hyaline ; hamuli 
five or six, stigma well developed, marginal cell open at apex, no submarginal cross 
veins in evidence, except sometimes a stump for the first. 

Some specimens show a very strongly reduced venation particularly of the 
posterior apical region of the forewing. The yellow rim to the compound eyes, so 
often noticeable in copal specimens, is believed to be due to the medium of their 

Remarks. The following are brief statements on the insect inclusions of each 
numbered piece of copal preserved in the British Museum collection. The first 
twelve pieces come from the Luke Thomas Collection. 

In.38982. One typical T. {Hypotrigona) gribodoi, and also a second inclusion of 
an insect believed to be a termite but whose head and thorax have been removed 
during the polishing of the copal. 

In. 38983. A well-preserved specimen of T. {H.) gribodoi, with quite a dark 

In.38984. Eight insect inclusions, i.e. six T. [H.) gribodoi, one dipteron and one 
termite. Some are most excellently preserved, and one of the bees shows an 
advanced reduction of the veins of the forewing. 

In.38985. One T. {H.) gribodoi and various other chitinous remains of insects, 
including aculeates. The bee is poorly preserved, portions of it having been worn 
away during the polishing of the copal, but sufficient remains for it to be identified. 

In.38986. Two T. {H.) gribodoi, both well preserved. 

In.38987. Copal too cracked for accurate diagnosis, but contains one insect 
which appears to be a T. (H.) gribodoi. 

In. 38988. A poorly preserved specimen of T. {H.) gribodoi. Copal around the 
insect somewhat cracked. A second inclusion, again rather poorly preserved, but 
markedly petiolate, is believed to be an ant of the genus Crematogaster. 

In. 38989. A T. {H.) gribodoi, well preserved. 


In. 38990. A well-preserved T. {H.) gribodoi, and a second unidentifiable inclusion. 

In. 38991. Two T. {H.) gribodoi, one very well preserved, the other somewhat 
less so, on account of its position on the edge of the piece of copal, which is slightly 

In. 38992. One well-preserved T. [H.) gribodoi. 

In. 38993. Two T. [H.) gribodoi, one better preserved than the other. Here the 
pubescence, so characteristic of the species, is plainly visible. Both show a little 
coating of hair. 

58511. All six insect inclusions appear to be T. {H.) gribodoi. In copal stated 
to be from Stettin on the Baltic, purchased from J. C. Rees 1867. There is, however, 
strong evidence from the so-called 'Baltic copal' from Stettin was imported from 
East Africa. 

In. 17686. Two T. {H.) gribodoi. Copal : East Africa. 

The characters of the following specimens agree more with the original description 
of T. bottegoi Magretti than with T. {H.) gribodoi Magretti. Unfortunately, our 
study of Recent and fossil material leads us to believe that the characters selected 
by Magretti are not constant enough for the separation of the two species, and we 
have thus been compelled to sink the former species under the latter.^ 

In. 18229. A well-preserved T. {H.) gribodoi with a detached head of the same 
species. The colour of the insect has been somewhat intensified by the copal and 
part of the ventral surface of the thorax, abdomen and legs appear whitish. This is 
probably because of air bubbles under the chitin of the insect. 

In.18214. At first glance the insect in this piece of copal appears different from 
the other specimens mentioned above, owing particularly to the yellow-brown 
abdomen, ruby eyes and the reddish-tinted cloud on the apical region of the fore- 
wings. This intensified colour, however, appears to be some effect of the copal. 

In. 18220. Two T. {H.) gribodoi, and additionally two well-preserved detached 
legs almost certainly belonging to the same species. One of the two insects is well 
preserved but the upper portion of the head and thorax has been pared away during 
the polishing of the copal and it is surrounded by cracks. The second insect is also 
well preserved but again has a distinct reddish tinge, particularly noticeable in the 

In. 18226. A single T. {H.) gribodoi, fairly well preserved but with the head 
partly pared away. Thorax and legs show many white 'air-bubbles'. On the abdo- 
men the pale bands indicate exposure of the thin intersegmental membrane as a 
result of distension of the abdomen. 

58622. A single specimen surrounded by fairly copious air-bubbles in the copal. 
However, from the side, the specimen is relatively clear to view and is seen to belong 
to T. {H.) gribodoi. A white rim to the eye and the white sheen to the wings are 

* The characters chosen by Magretti (1895 : 157) to separate T. bottegoi from T. gribodoi were: 'per la 
superfice minutissimamenta punteggiata, quasi Uscia e nettamente splendente del capo, del mesonoto e 
dello scudetto; per presentare I'area dell'ocello mediano e la linea mediana frontale leggiermente distinte 
ed inline per la colorazione giallo-testacea dello scapo antennale, delle mandibole, del labro et delle due 
paia di zampe anteriori.' [. . . in having the surface very finely punctured, being almost smooth and 
distinctly shiny on head, mesonotum and scutellum; in showing the area of the median ocellus and the 
median frontal line faintly distinct and finally by the yellow-testaceous colour of the antennal scape, of 
the mandibles, labrum and the two anterior pairs of legs.] 


believed to be spurious effects. It is slightly larger than the others, but the size of 
worker bees often varies considerably. 

Subgenus TETRAGONA Lepeletier & Serville 1828 
*i828 Tetragona Lepeletier & Serville : 710. 

Genotype. Trigona elongata Lepeletier & Serville 1828 (= Centris clavipes 
Fabricius 1804). 

Diagnosis. The inner face of the hind tibiae unevenly gabled but with a sharply 
elevated, median, plateau-like area that extends from the base almost to the apex 
and contrasts strongly with a wide, flat, posterior rim (particularly widened at the 
apex) ; the contour of the hind tibiae more or less clavate, with the apex in all cases 
much wider than the base and sometimes conspicuously expanded ; plumose hairs 
in addition to simple hairs usually present along the posterior lateral contour of the 
hind tibiae or present over their outer face near the base, or both. 

Remarks. The above is verbatim from Schwarz (1948). 

^1896 Meliponorytes succini 
1907 Meliponorytes succini 
I909d Meliponorytes succini 
1925 Meliponorytes succini 
1928 Meliponorytes succini 
1948 Meliponorytes succini 

Trigona (Tetragona) succini (Tosi 1896) 

Tosi : 352, pi. 6, figs 1-8. 

Tosi ; Handlirsch : 892. 

Tosi ; Cockerell : 5. 

Tosi ; Schroder : 254. 

Tosi ; Wheeler : 97. 

Tosi ; Schwarz : 8. 

1964 Meliponorytes succini Tosi ; Kerr & Maule : 2. 

HoLOTYPE AND Paratypes. Museo di Mineralogia deH'Univ. di Bologna. 

Distribution. Miocene, Sicilian amber. 

Diagnosis. Length 4 mm ; head subquadrangular, mandibles apparently with 
only two denticles ; inner surface of hind tibia plateau-like, of sharp elevation, with 
the posterior and apical rim fairly flat and wide ; inner face of hind basitarsus 
contains a smooth, bristleless area ; the hairs along the posterior edge of tibia appear 
to be plumose. 

Description. Dark, except abdomen which tends to be reddish. Compound 
eyes large, oval and naked. Ocelli situated in a slightly curved line, the lateral ones 
prominent, the median one lying in a furrow. Face slightly convex, pubescent, and 
the facial area itself ellipsoidal, slightly hairy and prominent. Clypeus almost 
triangular and provided with hairs. Vertex double-humped, divided by a median 
sulcus. Antenna twelve-segmented, the scape long, about one-third the length of 
the antenna, creased at point of insertion. The structure of the antenna is given in 
some detail by Tosi, but cannot be said to differ materially from the Recent Tetragona, 
unless Tosi's implication of the rather naked first to third flagellar segments is 
regarded as significant. Labrum extended, 'becoming round and hollowed like a 
bowl'. Mandibles with two denticles. Tongue hairy with dilated apex. Labial 


palps apparently four-segmented with long first segment. Thorax essentially as in 
Recent Tetragona, with long hairs on ridge of scutellum. Tegulae well developed. 

Wings hyaline but hairy, not extending much beyond the abdomen which appears 
to be fully extended (maybe distended). The main features of the forewing are as 
follows. Stigma very distinct, lanceolate in form ; first submarginal cross vein not 
strong ; marginal cell open at its extremity. 

An antennal cleaner is present, and its 'auschnitt' lined with hairs. Basitarsus 
as long as the remaining tarsal joints taken together and covered with setae. 
Posterior tibia well developed and covered apically with thick, long hairs, but basally 
with fewer and more robust hairs. Last tarsal joint larger than the others (except 
the basitarsus) and cylindrical. Pulvillus large. 

Abdomen ovoid, segments all the same size with longer hairs along posterior rim. 
First abdominal segment hollowed anteriorly. Ventral surface hulled and its first 
segment furnished with rows of longish hairs. A median sulcus follows the shape of 
the rows. 

Remarks. This species has been placed in the subgenus Tetragona not only on 
general morphology, but particularly on the structure of the hind leg. If Tosi's 
figures are carefully examined the inner surface of the hind tibia shows both a main 
ridge and also a clear dark zone posterior to it. This provides reasonable evidence 
that the inner surface is indeed plateau-like, of sharp elevation, with a fiat posterior 

Schwarz (1948) who reviewed Tosi's species came to no definite conclusion with 
regard to them. He did, however, raise some meticulous points, and rightly or 
wrongly insisted upon very minute and difficult detail. He refers, for instance, to 
the two denticles on the outer half of the apex of the mandible instead of the inner 
half. It is true Tosi illustrated them in such a position and number, and this may 
have been accurate for a Miocene genus, but it must in any case have been an 
achievement for Tosi to recognize denticles at all. The distinction of inner and 
outer halves may have been well-nigh impossible. Again the appearance of oblique- 
ness of the lateral ocelli is obviously influenced by the diffraction of light through 
the uneven amber surface and cannot be given the significance which Schwarz 
would like to attribute to it. The latter's reference to the length of the wing and 
to the stigma is non-committal. 

Trigona (Tetragona) iridipennis Smith 1854 
(PI. 2, figs 8, 9) 

*i854 Trigona iridipennis Smith : 413. 

1858 Trigona laeviceps Smith : 51. 

192 1 Meliponorytes (?) devictus Cockerell ; Cockerell : 544, text-fig. 4. 

1922 Trigona laeviceps Smith ; Cockerell : 714. 

1922 Meliponorytes (?) devictus Cockerell ; Cockerell : 714. 

1964 Trigona [Tetragonula) devictus Cockerell ; Kerr & Maule : 11. 

Specimens studied. British Museum (Natural History), In. 20702 (holotype of 
Meliponorytes (?) devictus Cockerell) and In. 43809 (identified by Cockerell as Trigona 
laeviceps Smith). 


Fossil distribution. Probably Pleistocene, Burmese amber : Hukong Valley. 

Diagnosis. Length 4 mm. Head and thorax black, face with short, silvery 
pubescence rather variably distributed, labrum and mandibles ferruginous, antennae 
testaceous, margins of thorax and scutellum fringed with fuscous pubescence and 
the sides of the metathorax with a silvery pubescence, elsewhere smooth and shiny ; 
wings subhyaline, iridescent ; abdomen ferruginous or the base testaceous, smooth 
and shining, extreme apex pale testaceous, ventral surface with short, silvery 
pubescence ; apical joints of tarsi testaceous. 

Description. Cockerell's description of specimen In. 20702 apparently reflects 
the distortion which would be encountered in looking through the spherical bead 
containing the specimen, but the bead has now been cut and the following details 
may be regarded as reliable. Length 4 mm. Head and thorax black, the abdomen 
and mandibles ferruginous, tarsi testaceous. The naked eyes are red but this cannot 
be considered the natural colour, the head is broad but not unnaturally so, and the 
ocelli are prominent, in a curve on the vertex, and have also taken on the reddish 
tinge. The antennae appear normal in position, size and shape for T. {T.) iridi- 
pennis ; the scape certainly cannot be considered long and the curve is only slight ; 
the flagellum is thick and short. 

The mesothorax is raised and the scutellum projects, but neither of these features 
is striking. Hairs are not much in evidence but a few can be perceived on the 
thorax and on the ridge of the scutellum ; this apparent nakedness, however, is 
what must be expected of white pubescence on fossilization in amber. 

Femora not especially robust, hind tibiae broadened at apex, triangular rather 
than clavate. Hind basitarsi are at most only half the length of the tibiae and 
cannot be described as large ; on the other hand, the abdomen is not short for the 
size of the creature. Forewings subhyaline with a well-developed pale stigma, but 
folded over the body so that the venation cannot be determined, except that the 
junction of the basal nerve and cross vein cu-a is coincident. Claws simple, pulvilli 

The specimen In. 43809 was referred by Cockerell to the Recent Trigona laeviceps 
Smith, which is synonymous with T. [Tetragona) iridipennis Smith. 

Remarks. Although the specimens look different, when examined superficially, 
the abdomen of 'M. devictus being long and that of T. laeviceps short, broad and 
curved downwards, they are conspecific since the differences in the abdomen are due 
to accidents of fossilization, the first being inflated by gases of putrefaction, the other 
strongly constricted, as if dried. 

T. laeviceps Smith had already been synonymized with T. iridipennis Smith by 
Schwarz (1948). On the other hand, the comparison of M. devictus with Recent 
species has led us to its identification with T. {T.) iridipennis. Cockerell, therefore, 
was mistaken in assigning these two specimens to two different genera and species. 

The reasons for Cockerell's mistake are interesting. In 1921 when Cockerell 
investigated the first piece of amber, he was under the impression that it was of 
Miocene age and in consequence placed his specimen under a Miocene genus, 
Meliponorytes. It was in the following year that he received a communication to 


the effect that this amber was, in fact, very much more recent. Consequently the 
specimen described in that year was referred to the modern Trigona laeviceps Smith 
which, as we have seen above, is a synonym of T. {Tetragona) iridipennis Smith. 
Unfortunately Cockerell never rechecked the specimen he designated as Meli- 
ponorytes (?) devictus and in consequence the latter name has persisted in the litera- 
ture until now. 

Subgenus indet. 
Trigona sicula (Tosi 1896) 

♦1896 Meliponovytes sicula Tosi : 355, fig. 9. 
1928 Meliponoryies sicula Tosi ; Wheeler : 97. 
1948 Meliponoryies sicula Tosi ; Schwarz : 9. 

HoLOTYPE AND Paratypes. Musco di Mineralogia deU'Univ. di Bologna. 
Distribution. Miocene : Sicilian amber. 

Diagnosis. Three times larger than T. [Tetragona) succini, i.e. about 12 mm long ; 
mesonotum a little less arc-shaped, hind basitarsus narrower and longer. Mandibles 
with two terminal notches ; pterostigma wanting. 

Description. The following characters are common with those of T. {Tetragona) 
succini : (i) large oval eyes, (2) situation of ocelli, (3) antennae, (4) labial palps, 
(5) claws and pulvillus, (6) hairy tongue, (7) prominent scutellem with long hair. 
The species appears to have been rather dark in coloration and the wing very similar 
to that of T. (T.) succini except for the lack of stigma. 

Remarks. The subgenus unfortunately cannot be determined. The specimens 
themselves are sufficiently like succini to be considered to belong to Trigona, although 
they lack the stigma, but their large size is a little perplexing. Again, the specimens 
are not well preserved, and in consequence inadequately described. Tosi wondered 
whether the species was not a Melipona, but in view of the present limited distribu- 
tion of the latter in South America, and the prevalent ideas of the phylogeny of this 
group, the latter suggestion is not a happy one. Only a re-examination of the type 
may shed light on the problem. The diagnosis given above is of necessity compara- 
tive, in view of Tosi's manner of description. Yet it must be remembered that the 
stigma illustrated by Tosi is much as in some Tetragona. Again, the comb and 
pollen rake of the hind tibia are notoriously difficult to detect in some fossil forms 
and are poorly developed in some Recent members of Tetragona. Failure to detect 
them, therefore, may be taken as an indication of their smallness, not of their 

The decision to include the present specimens in Tetragona was mainly the result 
of a study of their leg contours, as already mentioned. The wing venation, however, 
caused difficulty. According to Tosi vein Cug is redundant and vein Cu^ meets 
vein A at the distal and lower apex of cell 2nd Cu. Nevertheless, a slight furrow, 
which appears to exist in this vicinity in Recent forms, could, if it existed in fossil 
forms, have given rise to the latter illusion. Again, since the junction of the veins 


Cuj, A and Cu is unknown in Apidae, it is fairly obvious that a mistake has been 
made here. With regard to cross vein cu-a, not only does it lie along a rather 
abnormal slope but its relation to the basal vein is peculiar. Possibly in his figure 
Tosi has rather unwittingly exaggerated the junctions of the veins in this region in 
an effort to clarify their course. 

More than eight specimens of these fossil bees were found in one piece of amber. 

Tribe APINI Latreille 1802 

Description. This Old World tribe is unique among bees in the form of the male 
genitalia which, unfortunately, are as a rule not accessible in fossil forms. 

The Apini have the following characters in common with the Meliponini : scutellum 
raised, malar space short, ninth sternum modified. Forewing with first recurrent 
vein angulate or curved. Hindwing with jugal and anal lobes. 

The following features are diagnostic for Apini. Eyes densely hairy in Recent 
forms. Sting well developed. Either one outer tibial spur on hind leg or none ; 
hind basitarsus usually broad as tibia and always rectangular, with a depstum, or 
the latter broadened into an auricle ; basitarsal brush without the small bristleless 
or differentiated area ; claws cleft. 

Wings never extending much beyond the body. Distal cells of forewing always 
complete, marginal cell parallel-sided for a large part of its length in Recent species, 
but not parallel-sided and smaller and deeper in early fossil forms ; pterostigma 
becoming reduced, not tapering beyond base of vein r. 

Remarks. Most of the known fossil Apidae belong to this tribe, the most 
important and interesting of all as it embraces the honey bee Apis mellifera (p. 249). 
As a result, the evolution of this remarkable insect can be reconstructed in a detailed 

Genus ELECTRAPIS Cockerell 1908 

* 1908b Electrapis Cockerell : 326. 
igogd Electrapis Cockerell ; Cockerell : 7. 

Genotype. Apis meliponoides Buttel-Reepen 1906, by monotypy (Cockerell 

Diagnosis. Body morphology not very bomboid (p. 157) but wing venation 
retaining bomboid characters. Either a single outer hind tibial spur or none. While 
the body may have a variable quantity of hair, that on the eyes is scant or non- 

Description. Variable in size and shape ; from 4 mm to 14 mm in length. 
Normal disposition of tip of abdomen slightly ventral and proximal. 

Head. Moderately large and somewhat square ; tuft of hair on the vertex often 
present ; compound eyes moderately convex and ocelli prominent. Male has eyes 
far apart and the facial quadrangle is squarish. 


Mouth parts are well known only from Cockerell's descriptions, which may not 
apply to all species. They are as follows. 'Mandibles trmicate, much as in Apis, 
but the cutting edge with two notches ; labrum semicircular, tongue elongate and 
both it and the labial palpi essentially as in Apis ; spoon on end of tongue small, 
maxillary palpi very small, thick, apparently 2-jointed, malar space very short ; 
anterior end of clypeus straight'. The little that can be discerned in E. apoides 
Manning, however, suggests a single notch in the cutting edge of the mandibles, and 
an elongate 'tongue' with a rather large flabellum. 

Antennae moderately long, sockets large ; scape relatively long and curved, 
usually broadened distally and often articulating with a 'knob-shaped' pedicel or 
first joint of the fiagellum. Male antennae longer than female. 

Thorax variable ; (i) long and broad in proportion to the size of the insect, 
tapering sharply distally, very reminiscent of some Trigona ; (2) less broad and 
more slender, proportionately more reminiscent of Apis ; (3) more spherical in 
shape, usually well covered with hair, somewhat reminiscent of Bombus. Scutellum 
always distinct and often prominent. 

Front leg. Antenna cleaner always present but structurally slightly different 
from Recent Apis. 

Middle leg. Outer surface of tibia well provided with stout hairs. A single 
spur present at apex of tibia, stouter than that of hind leg. Basitarsus with long 
strong hairs. 

Hind leg. Tibia much longer than basitarsus. Posterior fringe of hairs on the 
tibia well developed and believed to contain some plumose hairs among the simple 
ones. Corbicula and pollen rake present. Sometimes a ridge covered with fine 
hairs occurs on the inner surface of the tibia. Basitarsus rectangular, with a brush 
and often a small differentiated area. 

Claws bifid. Pulvilli fairly large. 

Abdomen. Usually oval in shape and, at most, only moderately clothed with 
hair. In E. {Roussyana) palmnickenensis the hairiness of the sternites resembles an 
abdominal brush, usually as broad as the tibia or nearly so. The shape of the 
abdomen as figured by Buttel-Reepen (1906) may not be typical of the whole genus ; 
another known type has one edge convex, more like that of a recent Apis worker. 
The 'auschnitt' of the auricle between tibia and tarsus is deep and well formed, but 
there is only an incipient development of the depstum. A single outer spur is 
typically present. 

Forewing. Pterostigma variable in size and shape. Marginal cell moderately 
long and generally broader than that of Recent Apis, ending away from wing margin 
in a stub. Vein second r-m always curved, mostly S-shaped, but much more up- 
right than in Recent Apis and meeting M very slightly distad of the junction of M 
and second m-cu ; first r-m at right angles to M ; base veins of cells first R^ and 
first Rs form a Y with first submarginal cross vein ; the junction of first m-cu with 
M is angular as described, but not illustrated, by Cockerell (igogd). This junction 
is beyond half the wing length and thus much more distal than in Recent Apis. 


The basal vein meets at the base or just proximad of stigma and is gently S-shaped. 
It runs into Cu slightly proximad of the junction of the latter with cross vein cu-a, 
or the junctions here coincide. Lower section of basal vein (M) not less than twice 
the length of the upper section of basal vein (Rs). Vein Cua is fairly short. 

Hindwing. Hamuli variable (6-16, cf. Chalcobombus with 8-11). Cross vein 
cu-a oblique, the slope being posterior and basad, as in Recent Bombus and opposite 
in direction to that of Recent Apis ; M runs almost to wing margin ; cross vein 
r-m well defined, but only at most, slightly oblique, while Rs, like M, almost reaches 
wing margin. The general shape is more elongate than Recent Apis, and the wing 
possesses distinct and well-characterized anal and jugal lobes. 

Wings, in general, hyaline, fuliginous or fuscous and of moderate size, rarely 
longer than the body length though from their point of insertion on the thorax 
they often extend beyond the abdomen. Sometimes a fairly large, dark-coloured 
cloud (about one-twelfth total area) is present in the forewing. Most wings are 
hairy, but there is some doubt as to whether this applies to all species. 

Remarks. Most authors have held Electrapis to be directly ancestral to Recent 
Apis. This view is likely to be correct regarding only Electrapis s. str., here treated 
as a subgenus. The other species show clear signs of deviation. On the whole, 
however, one cannot separate these groups as distinct genera, as too many characters 
overlap. It is in fact impossible to give clear diagnoses of subgenera, and the three 
subgenera here distinguished are to that extent arbitrary. They are : 

(i) a group with Apis-\i\ie characters : Electrapis Cockerell igogd, s. str. ; 

(2) a group with Bomhus-\\\ie body and Apis-Y\k.e venation ; Protobombus Cockerell 
1908b ; 

(3) a Trigona-like group : Roussyana Manning i960. 

Cockerell (i909d) and Zeuner (1944, 1951) considered that the species of Electrapis 
were social insects. As evidence Cockerell pointed out the occurrence of many 
specimens crowded together in a small piece of amber {E. meliponoides) , whilst 
Zeuner considered the structure of the collecting apparatus. On other grounds too, 
this contention can be supported. [See also Appendix, p. 257.] 

Subgenus ELECTRAPIS CockereU 1908 

* 1908b Electrapis Cockerell : 326. 
igogd Electrapis Cockerell ; Cockerell : 7. 

Genotype. Apis meliponoides Buttel-Reepen 1906, by monotypy (Cockerell 

Electrapis (Electrapis) apoides Manning i960 

*i96o Electrapis {Electrapis) apoides Matining : 306, pi. 5, fig. i. 
1970a Electrapis apoides Manning ; Kelner-Pillault : 16, pi. 2, figs i, 2, text-fig. 5A. 

HoLOTYPE. No. 94 in the Bursey Collection (see p. 159), now in the Museum 
National d'Histoire Naturelle, Paris. 


Distribution. Eocene, Baltic amber. 

Diagnosis. Body 6-5 mm long, hairy. Forewing 6 mm. Hind tibia compara- 
tively short and stout, with a single, much reduced, spur ; corbicula exceptionally 
well developed with strong curved hairs on the posterior edge ; basitarsus square 
and flat, but with the posterior edge markedly convex. Wings fuliginous with a 
cloud apically ; pterostigma large ; eight hamuli. 

Description. Specimen well preserved, blackish. Total length 6-5 mm ; 
thorax 2-75 mm long and 2-25 mm wide ; abdomen 2-75 mm long and 2-5 mm wide ; 
forewing 6 mm ; hind tibia 275 mm. 

Proboscis extended for approximately the length of the head, provided with a 
large flagellum. Antennae very much like those of E. {Roussyana) proava (p. 236), 
with a similarly shaped, knob-like pedicel. 

The forelegs differ from those of other species of Electrapis in the shape of the 
antenna-cleaner. The zone of articulation between tibia and tarsus is greatly 
reduced, and the latter limbs are rounded, and not square, at their point of insertion. 
The clasp, too, is more highly developed, now having a fully shaped lobe and a 
distal tip projecting over and against the distal edge of the notch. The middle legs 
cannot be studied in detail, since they lie too much under the body of the insect. 
They are, however, well provided with hairs and appear to possess a single strong 

The hind legs are well proportioned, and not lanky as in E. {Roussyana) proava, 
and the hairs of the posterior margin of the corbicula are possibly plumose. On the 
apical margin of the tibia, posteriorly, a fine-haired rake is present, and the 'auschnitt' 
of the auricle is well developed. Moreover, the internal spur on the hind tibia is 
present and visible on the left leg of the specimen as viewed from behind. It is 
thicker and about one and a half times as long as the hairs of the anterior edge of 
the leg, but it is evidently in a state of reduction, being so small as hardly to serve 
any useful purpose. The basitarsus, although squarish, is elegantly shaped, with 
graceful curved outline, and not coarsely square as illustrated by Cockerell for 
Chalcobombus. Its inner surface is equipped with strong, almost bristle-like, hairs, 
which are arranged in rows as can be seen for those situated at the distal extremity. 

The wing venation does not differ materially from that of E. {R.) proava. Possibly 
the basal nerve of the forewing is slightly less curved, and the radius stronger. The 
hindwing is preserved in its entirety and exhibits eight hamuli, and well-developed 
jugal and anal lobes. Only one anal vein is known to be present, but the two lobes 
are separated by a deep cleft. A line that could be regarded as 4H is probably no 
more than a fault in the amber. Veins Rs and M continue almost to the wing 
margin, cross vein r-m slopes basad posteriorly, and cross vein cu-a exhibits a similar, 
but more oblique, slope. 

On the whole, the creature itself has the habitus of a small Apis. The head is 
well proportioned and neatly tapering, less square than some other species of 
Electrapis, and the thorax has neither excessive width nor length. The abdomen, 
however, is still rather small, though the habit of holding the tip ventrally, together 
with the presence of an air bubble on this particular specimen, rather prevents 


accurate observation of this point. The wings still project some distance beyond 
the apex of the abdomen. 

Remarks. The specimen was kindly lent by the late Mr Bursey of Surbiton, 
Surrey, to whom we acknowledge our thanks. 

Electrapis (Electrapis) meliponoides (Buttel-Reepen 1906) 
(PI. 3. figs 5. 6) 

*i9o6 Apis meliponoides Buttel-Reepen : 158. 
1907 Apis meliponoides Buttel-Reepen ; Handlirsch : 892. 
1908b Electrapis meliponoides (Buttel-Reepen) ; Cockerell : 326. 
I909d Electrapis meliponoides (Buttel-Reepen) ; Cockerell : fig. 4. 
19 1 5 Apis meliponoides Buttel-Reepen ; Buttel-Reepen : 10, fig. 3. 
1915 Electrapis meliponoides Butt. -Reap. ; Buttel-Reepen : 12. 
1928 Apis meliponoides Buttel-Reepen ; Wheeler : 98. 
1928 Electrapis meliponoides (Buttel-Reepen) ; Wheeler : 98. 
1 93 1 Apis meliponoides Buttel-Reepen ; Statz : 40. 
1931 Electrapis meliponoides Butt.-Reep. ; Statz : 41, figs 2, 3. 

HoLOTYPE AND Paratype. Two Specimens in the Westpreussisches Provinzial- 
Museum, Danzig. 

Material. Cockerell's specimens (i male, 7 females), Koenigsberg Collection. 
Also Nos. 238 and 461 in Scheele Collection, Geologisches Staatsinstitut, Hamburg. 
No. 238 is also Typen-Katalog des Geol. Staatsinstitut, No. 149. 

Distribution. Eocene, Baltic amber. 

Diagnosis. Body 14 mm. Bod}^ hairy but scant hair on eyes ; basitarsus 
comparatively long, broad and flat, posterior edge not convex ; wings fuliginous. 

Description. Cockerell's description can be summarized as follows (the measure- 
ments quoted in hundredths of a millimetre by Cockerell are to a degree of precision 
as unnecessary as it is misleading). Black, much erect fuscous hair on head and 
thorax. Tibia and tarsus with coarse fuliginous hair ; wing venation remarkable 
only for the small, piceous, stigma ; it is thought, too, that the upper section of the 
basal vein is comparatively short. Male specimen with eyes far apart ; facial 
quadrangle square ; antennae long, over 5 mm with last joint obliquely truncate. 

The following description is based on specimen No. 238 (Scheele Collection), which 
one of us (F. J. M.) was able to study in Hamburg. At first sight the fossil appears 
to be poorly preserved, in fact so much is covered with 'schimmeF that only a few 
segments of the abdomen can be discerned. The left fore wing and hindwing, how- 
ever, are both completely intact and the right forewing also shows much of the 
venation. Moreover, many parts of the legs are preserved, as well as the sting, and 
the hairiness of the thorax and the colour of the abdomen are clearly perceptible. 
Unfortunately, the head is completely missing. 

The specimen has the following measurements : overall length, without head, 
12 mm ; length of forewing, 9-5 mm ; hindwing 6 mm ; basitarsus 2-25 mm. 

The copious hairs on the thorax are erect and fuscous. The abdomen is black, 
but each segment appears to have a band of dark brown distally ; it is hairy but the 


hairs are smaller and darker than those on the thorax. They are, however, longer 
at the apex of the abdomen. The sting protrudes considerably and is strong and 
straight. In Recent Apis a straight sting is characteristic of the worker, that of the 
queen being distinctly curved. 

The many fragments of legs cannot be entirely pieced together. It is evident, 
however, that the claws are bifid and the distitarsi cordiform. The tibia, which 
can easily be recognized as such, has unfortunately had the outer surface pared away 
during the polishing of the amber. The corbicula, therefore, is lost, but the strong 
hairs of the anterior edge, which have not been destroyed, give complete proof of its 
earlier presence, if such were needed. There is no evidence for a tibial spur. This 
does not imply absence in view of the fragmentary character of the legs. The basi- 
tarsus is large, broad and flat, three-quarters the length of the tibia, the posterior 
edge quite straight and the posterior distal angle quite acute. Unfortunately the 
details of the structure at the junction of the basitarsus with the tibia are mostly 
masked by decayed pollen so that the depstum is not known. The strong non- 
plumose hairs of the anterior apex of the tibia, however, are present exactly as in 
Recent Apis and there is reason to suspect that the pollen rake has strong teeth. 
There is no evidence whatever for the presence of a pollen comb, but the pollen 
brush of the basitarsus is present. The strong hairs are arranged in rows with 
pollen still adhering to them. Approximately 17 hairs comprise the bottom 

The hairy forewing has a large cloud, and is slightly papillate apically. The 
wing venation is that of Electrapis but a slight elongation of the wing itself has 
brought about a few minor modifications. These are a narrowing of the apex of the 
marginal cell, an elongation of the third submarginal cell and the second M. Proxim- 
ally there is an increase in the length of the basal nerve with a consequent compen- 
sating alteration on the shapes of the first submarginal cell and the first M. The 
junction of basal nerve and cross vein cu-a is coincident on Cu, and vein cu-a slopes 
slightly apicad. Pterostigma piceous, but not small, though rather weak. 

The hindwing is particularly interesting because of the increase of the number of 
hamuli to 16. 

The other specimen from the Scheele Collection (No. 461) is poorly preserved, 
and covered with much 'schimmel'. Only part of a hindwing is well preserved. The 
insect is reminiscent of specimen No. 238, agreeing with it in all available characters. 
Length, with head, at least 13 mm ; hind basitarsus rectangular and long, almost as 
broad as tibia, and with a straight posterior edge ; hindwing in venation and shape 
agreeing with No. 238 except that the base is slightly shorter and the lobes deeper, 
and cross vein r-m appears more oblique. These characters can be accounted for 
by the oblique angle at which the wing must be viewed through the amber. At 
least 16 hamuli are present. 

The remaining features are taxonomically unimportant. They include the 
presence of two well-developed antennae inserted on the lower half of the face ; 
thorax possibly well supplied with hairs ; wings extending to tip of abdomen and 
probably beyond ; and the presence of legs whose broad outlines only can be 


Remarks. Cockerell defined the hind basitarsus of his specimens as 'broad and 
flat, as figured by Buttel-Reepen [Apis meliponoides)' . His specimens are thus 
considered to be identifiable with the latter. 

Buttel-Reepen was rather obsessed with the idea that E. meliponoides was inter- 
mediate in character between Recent stingless bees and Recent honey bees. What 
is probably true is that both groups evolved from some common ancestral form and 
that the Eocene E. meliponoides represents one member of this evolving series. 
Moreover, Buttel-Reepen stressed his point exclusively with regard to the hind 
basitarsus, which alone he described. From the descriptions here given, however, 
it is clear that the basitarsus has little that can be described as typically meliponid. 

Electrapis (Electrapis) tornquisti Cockerell 1908 

*i9o8b Electrapis (?) tornquisti Cockerell : 326. 
igogf Electrapis (?) tornquisti Cockerell ; Cockerell : 23. 
1928 Electrapis tornquisti Cockerell ; Wheeler : 98. 
193 1 Electrapis (?) tornquisti Cockerell ; Statz : 41. 
193 1 Electrapis tornquisti Cockerell ; Salt : 145. 

HOLOTYPE. Geological Museum, University of Koenigsberg. 

Distribution. Eocene, Baltic amber. 

Diagnosis. Length of body 11-5-12 mm ; robust and more Bombus-like than 
the previous species ; head and thorax abundantly hairy, but face with scant hair ; 
fourth antennal joint conspicuously shorter than fifth ; wing hyaline, stigma almost 
lacking ; hindwings with 15 hamuli ; abdomen broad. 

Description. A hairy bee, more so than E. {E.) meliponoides. Coloration of 
thorax anomalous. The broad abdomen, although apparently black, has the hind 
margins of the segments rather broadly seamed with white. Pubescence of abdomen 
short and pale. Tegulae large with very fine delicately plumose hairs. 

Venation differing from E. {E.) meliponoides in the rather distal position of the 
cross vein cu-a which lies slightly distad of the junction of the basal vein with Cu. 
It further differs in the lower part of this vein being strongly bent basad. It agrees 
with it, however, in that section Rs of the marginal cell is concave. 

Remarks. Although this specimen, on general morphology, is more bomboid 
than other Electrapis s. str., it has been retained in this section for convenience. 
The species was listed with some characters in 1908, but was not fully described 
until 1909. 

Subgenus PROTOBOMBUS CockereU 1908 

*i9o8b Protobombus Cockerell : 327. 
i909d Protobombus Cockerell ; Cockerell : 10. 

Genotype. Protobombus indecisus Cockerell 1908, by monotypy. 
Diagnosis. Wings comparatively apoid, and more bomboid, than in E. {Elec- 
trapis) . 


Description. Body globose, compact and Bombus-like. Compared with E. 
{Electrapis) it appears to have a greater range of notches on the mandibles (1-3), 
the elevation of the scutellum is accentuated and the inner tooth of the claws is 
shallower. It agrees with E. [Electrapis) in size (7-10 mm), in its hairless eyes, 
comparatively large wings, gently convex posterior margin of the basitarsus, tibial 
spur of the middle leg, large ocelli and large pulvillus. The hind tibial spur may or 
may not be present. 

Forewing. Stigma almost wanting. Marginal cell longer than in E. [Electrapis) 
but still ending obtusely at a distance from wing margin. All the submarginal cells 
exhibit peculiarities, in particular the second which is, according to Cockerell, 'very 
broad, pentagonal, much longer than third on marginal, and receiving ist recurrent 
nervure (ist m-cu) at or a little beyond end of its first third'. The latter feature is 
remarkable in that it shows a definite advance towards the Synapis wing. This 
point, however, is further emphasized by the first and second submarginal cross veins 
being 'equally oblique in opposite directions', and also by the fact that the third sub- 
marginal cell is produced apically, by implication presumably more so than in the 
E. [Electrapis) wing. In these points the wing is markedly apoid. It is not a 
Synapis wing, however, since the base of the third submarginal cell receives the 
second m-cu vein one-sixth from its end, and the first m-cu has a bend or angle in 
its upper part. Such characteristics are typical of the genus Electrapis as are also 
the remaining characters quoted by Cockerell. 

Hind wing. Vein cu-a strongly oblique with sometimes a slight double curve. 

Remarks. Cockerell considered this genus to be ancestral to Bomhus. This 
point of view, however, is difficult to maintain since Protobombus has a more apoid 
wing than even E. [Electrapis), though its body is somewhat reminiscent of Bombus. 
The wide divergence from Bombus may be more easily appreciated if the evolutionary 
sequence of the second submarginal cell, in relation to the position at which it receives 
the first m-cu vein, is summarized : 

Chalcobombus - beyond (distal of) the middle. 

Electrapis [Electrapis) - just beyond or at the middle, 

Electrapis [Protobombus) - much [Cockerell's term] before the middle. 

Synapis - about one-third before middle. 

Apis - about one-quarter before middle. 

Electrapis (Protobombus) indecisus (Cockerell 1908) 

*i9o8b Protobombus indecisus Cockerell : 326. 
igogd Protobombus indecisus Cockerell ; Cockerell : 10, fig. 5. 
193 1 Protobombus indecisus Cockerell ; Salt : 146. 

HoLOTYPE. Geological Museum, University of Koenigsberg. 
Distribution. Eocene, Baltic amber. 

Diagnosis. About 10 mm long ; mandibles obliquely truncate with three notches 
on cutting edge ; basitarsus broad. 


Description. A blackish bee, but base of abdomen brownish, with sparse pale 
hair. Thorax and occiput densely covered with rather short, plumose, whitish 
hair. Legs and tegulae on the whole blackish with pale hair. The distinctive 
feature of the pale fuliginous wing is a dark cloud at end of marginal cell. 

Remarks. Listed by Cockerell with a few characters in 1908, but not fully 
described until 1909. 

Electrapis (Protobombus) tristellus (Cockerell 1909) 

*i909f Protobombus tristellus Cockerell -.24. 
1931 Protobombus tristellus Cockerell ; Salt : 146. 

HOLOTYPE. Geological Museum, University of Koenigsberg. 

Distribution. Eocene, Baltic amber. 

Diagnosis. About 7 mm long ; much erect hair on vertex ; wings approximately 
67 mm long ; abdominal segments black with hind margins 'rather broadly pale 
reddish' ; anterior margin of basitarsus gently convex ; glossa, as preserved, short ; 
mandibles long, falciform with a single deep notch ; last two joints of labial palpi 
rather large ; eyes wide apart above. 

Description. A blackish bee ; mesothorax and scutellum as well as vertex with 
much erect black hair, abdomen as described above. The forewing has a large, 
conspicuous fuliginous cloud. Venation as in E. {Protobombus) indecisus, except 
for a presumed double curve of the long, oblique vein cu-a of hindwing. Mouth 
parts appear to be well preserved and differ in several respects from those of E. 
(Electrapis) meliponoides , for instance in the short broad glossa and the large 
flabellum. The length of the mouth parts, however, is difficult to ascertain in fossil 
forms, since they may not be completely extruded. Little reliance, therefore, should 
be placed on this character for diagnostic purposes. 

Remarks. This specimen was not listed by Cockerell in 1908. 

Subgenus ROUSSYANA Manning i960 
Genotype. Apis palmnickenensis Roussy 1927, by original designation. 
Diagnosis. This subgenus embraces the more Trigona-like members of the genus 
Electrapis. These bees exhibit, however, all the characters of this genus, in so far 
as they are known. They are distinguished by the habitus of the body which is 
reminiscent of the stingless bees. 

Electrapis (Roussyana) palmnickenensis (Roussy 1937) 
(PI. 3, figs I, 2) 

*i937 Apis palmnickenensis Roussy : 66, fig. 13. 
1938 Apis palmnickenensis Roussy ; Armbruster : 89. 
i960 Electrapis {Roussyana) palmnickenensis (Roussy) ; Manning : 306. 

HoLOTYPE AND Paratypes. Roussy Collection, Aigle, Switzerland. 


Material. In addition to the types, one specimen in Geologisches Staatsinstitut 
Hamburg (Scheele Collection, No. 218 ; also listed in Typen-Katalog des Geol. 
Staatsinstitut, No. 150). 

Distribution. Eocene, Baltic amber. 

Diagnosis. Small, black insects about 4 mm long ; scutellum very hairy ; 
sternites with rows of hairs. Inner surface of tibia with a ridge of short hairs cover- 
ing about a quarter of the posterior surface. Basitarsus less than half the length of, 
and narrower than, the tibia, its posterior upper apex exhibiting an area clear of 
bristles and an incipient depstum. 

Description. We have inspected M. Roussy's piece of amber containing eight 
specimens, two of which are complete. The following is the condition of the 
individual specimens, their numbers agreeing with the semicircular alignment from 
the left upper corner of the piece of amber towards the right lower corner. 

I. Badly preserved, except fore and hind legs. Wing venation masked by 

'schimmel'. Possibly not more than 3-5 mm long. 
2 and 3. Very much alike and well preserved ; 3-4 mm long. Details of almost 

all appendages can be discerned including mandible and proboscis of No. 3. 

Spur on hind legs distinct. Wings of No. 2 not clear. 

4. The best preserved specimen, but parts of some legs cut off. A considerable 
amount of wing venation can be discerned. 3-5 mm long. 

5. Much covered by 'schimmel' but much of the wing venation recognizable. 
Antennae prominent. 

6. Badly preserved and consists only of remains of thorax and legs. 

7. Somewhat covered with 'schimmel' and wings cut off short. Insect bent on 
itself. One antenna present and well preserved. 

8. Remains of a head with mandible. 

The wings extend well beyond the apex of the abdomen ; the head is broad, but 
less so than the thorax. The antennae are inserted on the lower part of the face 
and consist of a comparatively long, slightly curved scape and 11 segments to the 
fiagellum ; the pedicel appears to be knob-shaped. The eyes are prominent on the 
side of the head, and the ocelli distinct. The thorax is strong and carries a prominent 
scutellum. The abdomen is short and oval. The forelegs exhibit a well-developed 
antenna cleaner with broad flap and deeply rounded 'auschnitt', whilst the hind leg 
is characterized by a distinct corbicula with scopa, strong pollen rake, and short 
outer tibial spur but no pollen comb. The basitarsus is rectangular and equipped 
with a hairy pollen brush on the inside. 

In addition to the eight specimens of M. Roussy, another has been found. It is 
preserved in the Scheele Collection, Hamburg. Through the kindness of Professor 
E. Voigt and Dr Walter Hantzschel it was lent to us for detailed inspection. Since 
the piece of amber contains one complete insect, perfectly preserved, and two frag- 
ments of legs, one in excellent condition showing tibia and tarsi, it has been possible 
to obtain a reasonably complete description of it, and the following is based on this 


Size. From head to tip of wing 4-25 mm. Length of forewing 3-25 mm, 
extending 1-25 mm beyond the tip of the abdomen. Length of hind tibia and tarsus 
2 mm. This specimen appears to be longer than the Roussy specimens, but this is 
deceptive. Though the wings extend conspicuously beyond the abdomen, the length 
of the body (which cannot be measured accurately) is possibly no more than 3 mm. 

Colour. Very dark brown, or blackish. A creamy shading on the notum may 
be due to the natural colour of its copious hairs, or else it is due to the presence of 
some pollen. 

Head moderately large, black, face sparsely hairy, with cream-coloured pubescence 
on the vertex. There is a reddish patch around the right ocellus, but this is almost 
certainly the result of post mortem alteration. 

The compound eyes are hairless and prominent, and almost reach the bases of the 
mandibles, so that the malar spaces are much reduced. The three ocelli are large, 
set on the vertex and arranged in a shallow triangle. The single median frontal 
suture runs from the middle ocellus downwards. 

The clypeus is trapezoid, tapering dorsally because of the convergent slope of the 
lateral epistomal sutures, and it is not protruding. The mandibles appear to have 
at least a convex cutting edge. The anterior part of the face terminates in a narrow 

The antennae are not more than moderately long, inserted close together im- 
mediately above the frontoclypeal suture and apparently yellow and black. The 
scape has slightly less than half the length of the flagellum. The flagellum consists 
of eleven segments mostly subequal in size, though the terminal segment is longer, 
and those near the pedicel thinner and more tapering, and also darker in colour. 

Thorax. Well developed ; pronotum short ; scutellum very prominent. The 
whole thorax copiously covered with hairs which are longest on the scutellum, but 
shorter on the pronotum. Tegulae large. 

Abdomen. Small, oval. Tergites not noticeably hairy, though the sternites 
have rows of well-defined hairs. No intersegmental membranes can be discerned 
and this is taken as evidence of the rather contracted nature of the abdomen. 

Wings. Hairy and fuscous ; pattern of wing venation as in Electrapis, with the 
following points of interest ; in the forewing the pterostigma is large and deep ; 
the anterior vein bounding the marginal cell, i.e. radius, is not coincident with the 
wing margin and its course continues beyond the apex of the marginal cell towards 
the wing apex ; cross vein cu-a lies distad of junction of basal nerve with Cu ; the 
hindwing has only six hamuli and both jugal and anal lobes are well developed. 

Three anal veins are present in the hindwing. They can be seen if the left hind- 
wing is viewed head-on, i.e. as inspecting the face of the insect. 

Legs. Antenna cleaner present on foreleg, 'auschnitt' large, but since the legs 
are flexed on themselves other structures cannot be clearly identified ; middle leg 
has tibia at least as long as femur, broadening towards apex, and covered with strong 
hairs, particularly posteriorly. Pollen is entrapped in these hairs. A distinct tibial 
spur can be recognized on this leg. 


The hind leg appears to have a comparatively large trochanter distinctly tapering 
towards the base ; tibia longer than femur, triangular in shape, and with a well- 
developed scopa ; hairs appear somewhat plumose ; corbicula moderately well 
developed, smooth, with few hairs within the corbicula itself ; inner surface of tibia 
with the ridge of short hairs occupying such a position posteriorly that the rim which 
separates it from the posterior edge of the corbicula is very narrow but the anterior 
rim is wide. The tibial appendages consist of a small, outer tibial spur, straight, 
and non-serrate, and a strong pollen rake projecting on a shelf from the inner apex 
of the tibia ; no pollen comb is present, but at the posterior apex of the tibia an 
angle suggests an incipient auricle. The tibia is not strongly curved, even pos- 
teriorly, but the isolated hind leg, because of the curve of the amber, deceptively 
gives this impression. 

The basitarsus is narrower than the tibia and rectangular. The two posterior 
apices are strongly acute ; the inner surface consists of a brush of strong hairs, whose 
disposition in rows is rather masked by the comparatively great length of the bristles ; 
possibly five rows is the maximum, though the bristles per row cannot be computed ; 
the 'clear' area is present around the posterior upper apex. Distitarsi cordiform. 

The claws appear entire at first sight but close inspection reveals a single tooth at 
about half-way down. Pul villus well developed. 

Remarks. In 1938 Armbruster criticized Roussy's description of these insects, 
which admittedly was in a popular style. He failed, however, to appreciate their 
significance since he was unacquainted with the Trigona-like group of Electrapis. 
The very detailed description here given will, it is hoped, fill an awkward gap and 
establish this group as a natural division of the primitive honey bees. (See also 
p. 254.) 

Electrapis (Roussyana) proava (Menge 1856) 
(PI. 3, figs 3, 4) 
*i856 Apis proava Menge : 26. 
1891 Apis proava Menge ; Scudder : 689 (5938). 
1907 Apis proava Menge ; Handlirsch : 892. 
igogd Apis proava Menge ; Cockerell : 7. 
1915 Apis proava Menge ; Buttel-Reepen : 10. 
1928 Apis proava Menge ; Wheeler : 98. 
193 1 Apis proava Menge ; Statz : 40. 
1931 Apis proava Menge ; Salt : 143. 

Lectotype. British Museum (Natural History), In. 43592, herein selected. 

Paralectotype. British Museum (Natural History), In. 18757. 

Distribution. Eocene, Baltic amber. 

Diagnosis. Body about 6 mm long (after some allowance has been made for a 
contracted and bent abdomen), length of forewing 6-5 mm. Scant hair on body 
as well as on eyes ; hind leg rather lanky, due to a proportionately long tibia, 
basitarsus broad, fiat and rectangular, posterior edge convex ; wings hyaline, almost 


Description. The lectotype, on which the following description is based, is well 
preserved, blackish in colour, with the terminal portion of thorax becoming paler, 
the whole body being very scantily supplied with hairs. The actual length of the 
body is 6 mm, length of abdomen approx. 2-25 mm, width of abdomen 2-25 mm ; 
length of thorax approx. 275 mm, width of thorax 2-5 mm ; length of hind tibia 
3-25 mm. 

Head squarish, poorly supplied with hair ; ocelli prominent. Mouth parts not 
known. Antennae as recent Apis, but pedicel 'ball-shaped' and scape broadened 
distally, meeting the pedicel steeply. 

Thorax proportionately very broad and thick, tapering sharply distally, not hairy, 
reminiscent of some species of Trigona. 

The foreleg has an antenna cleaner of a somewhat primitive type, and well 
characterized by the clasp, which is long and thin and completely overlaps the hollow. 
The zone of articulation of tibia and tarsus is greatly extended. 

The middle legs have a pronounced spur on the apex of the tibia. This spur is 
longer and sturdier than the corresponding spur of the hind leg. The outer edge of 
the tibia is well provided with hairs. 

Hind legs fuscous, a little disproportionate on account of the great length of the 
tibia, the latter provided with long, somewhat plumose, hairs along the posterior 
edge and a neat row of short hairs down the inner median line. A single spur is 
present but its position is such that it is difficult to see on the specimen. Basitarsus 
rectangular, with convex posterior edge, covered with hairs which are strong on the 
inner side. An auricle is possibly not present but a concavity is present between 
tibia and basitarsus. One gains the impression that this zone is slightly more 
primitive than in Electrapis {R.) palmnickenensis. There is a tibial rake, which is 
stronger and coarser than that of the latter species. 

The venation of the fore wing is characterized as follows. Stigma comparatively 
large, vein cu-a at right angles on A and meeting Cu slightly distally of the junction 
with the basal vein. Upper section of basal vein (Rs) apparently proportionately 
longer than that of E. meliponoides. 

Hindwing incompletely preserved. Eight hamuli, compared with six in E. [R.) 
palmnickenensis. One might be inclined to regard this as a meliponid feature, since 
E. {E.) tornquisti has 15, E. {E.) meliponoides 16 (though £". {E.) apoides has only 8), 
Apis {Synapis) henshawi has 17, Recent A. mellifera 20-23 and A. dorsata 23, but 
the smaller A.florea only 12. Recent Bombus species have 20-23 and Euglossa 27. 
There appears, however, to exist a relation between the number of hamuli, on the 
one hand, and body size and power of flight, on the other. 

The part of the hindwing which is preserved includes cu-a, r-m and sections of M 
and Rs, together with cell Cu. It agrees with E. {E.) apoides. 

The second specimen. In. 18757, is covered with 'schimmel', and is assigned to 
E. (R.) proava with some reservation. It is larger than the lectotype, being 8-i mm 
long. The forewing is 6-8 mm long and shows a dark apical cloud covering about 
one-twelfth of its area. The venation, in so far as it can be ascertained, appears to 
be identical with that of the lectotype. We hold that this is Menge's other specimen, 
as will be shown under 'Remarks'. 


Remarks. It appears that Cockerell (igogd) thought Httle of Menge's detailed 
description, and that he never saw the specimens, presumably because he expected 
them to be in the Koenigsberg Collection. 

We have satisfied ourselves that the two specimens, In. 43592 and In. 18757 of the 
British Museum (Natural History), are in fact Menge's types, though no specific 
identification was attached to them. Our reasons are as follows. Two specimens 
were available to Menge, one almost entire and the other covered with 'schimmel', 
i.e. with the products of the gases of putrefaction. The first lends itself to descrip- 
tion, but the second is in a very poor condition. Menge's description of the first 
tallies in a remarkable way with the British Museum specimen In. 43592, and in a 
negative manner so does the second, In. 18757. Furthermore, the history of the 
specimens supports this view. They were purchased in 1892 by the Trustees of the 
British Museum from Messrs Stantien & Becker, first placed in the Zoology Depart- 
ment and later (about 1904) transferred to the Department of Geology (now 

Fossil insects preserved in amber were sold by the above-mentioned firm over a 
long period of time. Their stocks were eventually taken over by the Prussian State 
and became the care of the State Amber Collection in Koenigsberg. Before this 
happened, however, Dr Richard Klebs, a geologist in the Prussian State Geological 
Survey, was in charge of the amber collection of Messrs Stantien & Becker. A set of 
346 specimens were purchased by the British Museum in 1892, which included the 
two bees in question. These specimens bear labels marked 'R. Klebs, Museum 
Stantien & Becker'. This suggests that they did not come directly from the amber 
works of the firm, but were acquired through Dr Klebs. There is other evidence 
that not only the Menge Collection, but also part of the Germar and Berendt Collec- 
tions was dispersed with the aid of this firm, and it would appear that Dr Klebs 
bought up such collections in the first instance and handed them over to the firm at 
a later date. For this reason specimens of such provenance are usually assigned to 
the 'Klebs Collection'. 

If the specimens In. 43592 and In. 18757 were obtained by the British Museum in 
1892 from Menge's collection via Dr Klebs and Messrs Stantien & Becker it is evident 
that Cockerell could not have found them in the Koenigsberg Collection in 1908. 

It now remains to be shown that Menge's descriptions indeed tally with those 
given here as the result of independent study. Of the well-preserved specimen, 
Menge says that it is 7 mm long, with the wings 10 mm, breadth 3-4 mm. The 
abdomen is bent so that the sting lies under the apex of the tarsi of the first pair 
of legs. The tarsus of the foreleg comprises an antenna cleaner, or rather a sinus 
('ausschnitt'). A spur is present on the tibia of the hind leg where it projects from 
its outer surface, and bristles are present on the digits of the hind feet. The bristles 
are a little irregularly shaped. The body itself is scantily haired and the ocelli and 
compound eyes are bare. The latter are oblong and narrow, and separated from 
each other. Twelve antennal segments are present, and these, together with the 
shape of the eyes, indicate for Menge a worker bee. 

This description is at variance with our observations in two respects. First, the 
number of segments of the antennae of the fossil cannot be ascertained. Those of 


the good specimen (and the other is too obscured by 'schimmel' for any rehable 
observation) has one flagellum cut off short at the surface of the amber, the other 
twisted into a difficult position. Menge, however, referred to twelve visible segments 
for the worker.^ The explanation of the discrepancy between descriptions and fossil 
is believed to be the treatment of the specimen whilst in the hands of Messrs Stantien 
& Becker. The amber pieces were reduced in size to the minimum, preferably made 
rectangular, and embedded in a resin in a glass cell on a microscope slide. The 
polishing process resulted in the loss of the distal portion of the antenna, and parts 
of the legs. Menge, therefore, may well have been able to study the entire antenna. 

The second particular in which the description differs from the fossil is Menge's 
reference to the abdomen being so bent as to lie under the tarsi of the front pair of 
legs. The abdomen of the fossil does appear to be bent under the body but certainly 
not to such a forward position as Menge suggested. Also no sting can be seen. 
The abdomen, however, does touch the tarsus of a leg which must be one of the middle 
pair, for both fore and both hind legs can be made out separately. But the dis- 
position of the abdomen is so obscured by the poor state of the amber in this region 
that it is possible for Menge to have made the above statement rather from con- 
jecture than from actual observation, especially as he would have expected the 
abdomen to be longer than it actually is, believing the species to be closely related 
to Apis mellifera. 

Though Menge was right in saying that the general morphology of the specimens 
resembles that of the honey bee in many respects, closer study has now revealed 
some features reminiscent of Trigona, such as the proportionately broad thorax, the 
short and round abdomen, and the small number of hamuli. 

Genus APIS Linneaus 1758 

*i758 Apis Linnaeus : 343, 575. 
1810 Apis Latreille : 439. 
1938 Hauffapis Armbruster : 43. 

Genotype. Apis mellifera Linnaeus 1758 (= Apis mellifica Linnaeus 1766), 
designated by Latreille (1810 : 439). 

Diagnosis. Medium-sized bees, with hair. Eyes usually hairy, clypeus not 
protuberant ; well-developed hind tibia, but spurs and comb lacking ; basitarsus 
rectangular and covered with bristles arranged in rows ; no reduction in wing 
venation, marginal cell always complete. 

Remarks. This genus comprises the Oligocene, Miocene and Recent Apini. They 
are all extremely closely related to each other, and the division into two subgenera 
is based on very unsatisfactory characters. Similarly, the numerous species and 
subspecies that have been named cannot all be maintained if the standard of Recent 
taxonomy is applied. 

9 This number is incorrect of course, for Recent Apis workers, if it refers to the flagellar segments, 
since it is the drone that has twelve and the queen and worker (females) eleven segments, but it was 
probably computed to include the scape. 


Subgenus SYNAPIS Cockerell 1907 

*i907 Apis {Synapis) Cockerell : 229. 
1931 Synapis Cockerell ; Statz : 42. 

Genotype. Synapis henshawi Cockerell 1907, by monotypy. 

Diagnosis. General morphology very much like that of Recent Apis but differing 
in the following points : labrum less tongue-shaped ; third segment of fiagellum 
short but broad ; basitarsus, though variable in shape, never identical with Recent 
Apis ; junction of veins cu-a, Cu and the basal vein coincident, or vein cu-a lies in a 
slightly more proximal position. 

Remarks. This group of honey bees has been extensively treated by Statz in 
1931 and 1934, who regarded it as a genus. Cockerell, however, did not grant it 
more than subgeneric status, regarding the venation characters as important. 
Cockerell's view has been adopted by the present writers. 

Apis (Synapis) cuenoti Theobald 1937 
*i937 Apis cuenoti Theobald : 401 ; pi. 8, fig. 16, pi. 28, fig. 8. 

HoLOTYPE. Ecole des Eaux et Forets, Nancy, No. F.173 (Coll. Fhche). 

Distribution. Oligocene (Stampian), Cereste, France. 

Diagnosis. Length of body 1375 mm ; eyes pubescent ; hair on body fairly 
plentiful ; second m-cu meets M near distal corner of cell second Rs ; forewing 
length 12-5 mm. 

Remarks. Not so well preserved as some other representatives of Synapis. 
Specific rank has been retained for this specimen since it is most difficult to reconcile 
the wealth of hair, the pubescence of the eye (compare A. (5.) henshawi kaschkei, 
p. 243, the extraordinary length of the wing and the distal juncture of second m-cu 
with M, with any other Synapis. Theobald compared it with Apis oligocenica (recte 
A. (5.) henshawi), and suggested some connection between the two. It is thus likely 
that upon re-examination the specimen may turn out to resemble this species more 
closely than is at present apparent. 

Apis (Synapis) henshawi Cockerell 1907 

♦1907 Apis [Synapis] henshawi Cockerell : 229. 

1907 Apis {Synapis) henshawi Cockerell ; Handlirsch : 1357. 

1915 Apis oligocenica Meunier : 210, pi. 21, fig. 4. 

19 1 5 Apis henshawi Cockerell ; Buttel-Reepen •.14. 

193 1 Apis {Synapis) henshawi Cockerell ; Statz : 42. 

1934 Synapis henshawi Cockerell ; Statz : 3, 6, tf. i, 4, 5, 6. 

1934 Apis oligocaenica Meunier ; Statz : i. 

1938 Apis oligocaenica Meunier ; Armbruster : 88. 

HoLOTYPE. Museum of Comparative Zoology, Harvard University. 
Distribution. Oligocene (Aquitanian), Rott, West Germany. 


Diagnosis. Body 12-16 mm, forewing 8-5-10 mm. Variably hairy, eyes naked. 
Second m-cu meets base of cell second Rs on the abscissa of vein M beyond half- 

Description. The three subspecies included in this species vary in size, the body 
being 12-16 mm. A. (5.) cuenoti would be included in this on the grounds of size, 
but its forewing is longer than in any known specimen of yl. (5.) henshawi. The 
hairiness varies, and the hairs are perhaps sometimes plumose on the vertex. These 
bees closely resemble A . mellifera in general appearance. 

Remarks. The first description of this species is by Cockerell (1907). He 
examined three specimens from Rott which the Museum of Comparative Zoology, 
Harvard, had obtained from the firm of Dr Krantz, Bonn, under the name of Apis 
dormitans Hey den 1862. One of these specimens was named Synapis henshawi by 
Cockerell, but since the other two bore Apis dormitans labels, he assumed that they 
belonged to Hey den's species, which Cockerell regarded as different from his 5. 

Curiously enough, Heyden also had used three specimens of the Krantz Collection 
for his description of Apis dormitans, so that it seemed possible that Heyden's speci- 
mens were those that were eventually sold to Harvard University. That this is 
not so, however, is revealed by the fact that Cockerell was able to describe wings 
of all three Harvard specimens, whilst Heyden stated that one of his specimens was 
'without visible signs of wings'. Moreover, the holotype of Apis dormitans Heyden 
is in the collection of the British Museum (Natural History), and its re-examination 
(p. 251) has confirmed the view held by other authors that Heyden's species is not 
an Apis. 

Apis dormitans Cockerell {nee Heyden), based on two specimens in the Harvard 
Collection, is a true Apis {Synapis), as was demonstrated by Statz (1931). Hence it 
becomes necessary to rename this form, the species name being preoccupied by 
Heyden's form. 

Three different Synapis have been described from Rott. They all come from the 
same beds and the same biotope. In view of the variation known to exist in large 
Apis populations, they are here reduced to subspecific status. 

Apis {Synapis) henshawi dormiens subsp. nov. 

(PI. 4, figs 1-5) 

*igo'j Apis dormitans [nee Heyden) ; Cockerell : 228. 

1908 Apis dormitans Cockerell ; Handlirsch : 1357. 

193 1 Apis oligocenica Meunier ; Statz : 46 (erroneously placed here). 

193 1 Synapis dormitans (Cockerell) ; Statz : 42, 46, figs la, 2b, 3c, 4a, 5-9. 

1934 Synapis dormitans (Cockerell) 

1944 Synapis dormitans (Cockerell) 

Holotype. Museum of Comparative Zoology, Harvard University. (Cockerell's 
first specimen, 1907, described as Apis dormitans.) 

Paratype. Second specimen ; Museum of Comparative Zoology, Harvard. 



46, figs 




figs 2, 8, 




figs I, 3 



Material studied. In addition to the holotype and paratype, two specimens 
in the British Museum (Natural History), Nos 59634 and In. 36655 ; also all the 
specimens in the collection of Dr Statz, Cologne. 

Distribution. Oligocene (Aquitanian) : Rott. 

Diagnosis. Body 13 mm, forewing 8-5 mm. Hair on vertex not plumose, eyes 
naked. Basitarsus with 9 rows of bristles, middle row numbering 24. Vein cu-a 
a little proximad of junction of basal vein with Cu. 

Description. The medium-sized form of the species. In addition to the 
characters stated in the diagnosis, the following are worth noting : the flagellum is 
not narrowed at the base, the basitarsus is i-6 mm long and 0-9 mm wide and the 
depstum only moderately developed. 

Remarks. Numerous other specimens exist, mostly collected by Dr Statz, to 
whom we owe the detailed description and excellent reproduction of specimens. In 
1931 Statz thought that Meunier's Apis oligocenica was synonymous with his Synapis 
dormitans, but after the study of Meunier's type he correctly synonymized it with 
5. henshawi (q.v.). 

Apis (Synapis) henshawi henshawi Cockerell 1907 

*I907 Apis [Synapis) henshawi Cockerell : 229. 

1915 Apis oligocenica Meunier : 210. 

1934 Synapis henshawi Cockerell ; Statz : 5, figs i, 4, 5, 6, 7. 

1944 Synapis henshawi Cockerell ; Statz : 63, figs 2, 5. 

Holotype. Museum of Comparative Zoology, Harvard University. 
Other material. Holotype of Apis oligocenica Meunier, Heimatmuseum, 
Siegburg an der Lahn. Five specimens in the Statz Collection. 

Distribution. Oligocene (Aquitanian) : Rott. 

Diagnosis. Body 15-16 mm, forewing 10 mm. Hair on vertex plumose or 
simple, eyes naked. Basitarsus with 10 rows of bristles (second very short), middle 
row numbering about 28. Vein cu-a coincident with junction of basal vein with Cu. 

Description. This is the largest of the three subspecies. Making allowance for 
the distended abdomen (measured length 16-5 mm), Cockerell inferred a length of 
15 or 16 mm. When Statz encountered the holotype of Apis oligocenica Meunier 
in the Siegberg Museum, he satisfied himself that it was conspecific with A. (5.) 
henshawi, and he described it in detail under that name in 1934. There appears 
indeed to be a close agreement between Cockerell's and Meunier's types, except that 
Statz states positively that the hair on the vertex is not plumose, whilst in Cockerell's 
specimen 'this hair appears to be plumose'. Evidently, Cockerell did not feel quite 
sure, and since the hair is simple in all other Synapis, he may have misinterpreted 
his specimen. 

The wings are strikingly short for the size of the body. The antenna cleaner is 
deeply incised, semicircular, and thus differs from the shallower type present in 
A. (5.) dormiens. The basitarsus has ten rows of bristles, whilst in A. (5.) dormiens 


only nine appear to be present, and the number of bristles comprising the middle 
rows is slightly larger. 

Apis (Synapis) henshawi kaschkei (Statz 1931) 

*i93i Synapis kaschkei Statz : 50, figs 10, 11. 
1934 Synapis kaschkei Statz ; Statz : 3, figs 3, 9. 

HoLOTYPE. In Statz Collection, Cologne. 
Distribution. Oligocene (Aquitanian) : Rott. 

Diagnosis. Body 11 -12 mm, forewing 9-25 mm. Basitarsus broad, and shorter 
than in any known Apis. Vein cu-a very proximad of junction of basal vein with Cu. 

Description. This is the smallest bee from Rott. It differs from the other two 
Synapis in the structure of the basitarsus which is very short and almost square 
(i-2 mm long and i-o mm broad). Statz, in saying that this basitarsus almost agrees 
with that of Apis iridiae Friese from Sumatra, is, however, mistaken, as this species 
has a pronouncedly oblong basitarsus. The depstum closely resembles that of 
Recent Apis mellifera Linn., but the wing venation places this form in the genus 
Synapis. There appear to be nine rows of bristles on the basitarsus, including the 
second row which is very short. It has a smaller body and a longer forewing than 
the other two Synapis. 

Remarks. Only three specimens are known, two of which, including the holotype, 
have been seen by one of us. Its status may have to be reconsidered when more 
material becomes known. For the time being, Statz's description cannot be im- 
proved upon. He figures the holotype (1931 : figs 10, 11) and a second specimen 
in his own collection (1934 : figs 3, 9), the latter figure being the diagram of the fore- 
wing venation derived from the two fore wings of this specimen. 

Subgenus APIS Linnaeus 1758 

*i758 Apis Linnaeus : 343, 375. 
1938 Hauffapis Armbruster : 43 (Genotype : Haujfapis scheuthlei Armbruster). 

Genotype. Apis mellifera Linnaeus 1758 (= Apis mellifica Linnaeus 1766), 
designated by Latreille (1810 : 439). 

Diagnosis. Third segment of flagellum shorter, but not broader, than the remain- 
ing segments ; the broadly rectangular, neatly curved basitarsus always possesses a 
distinct auricle ; cross vein cu-a well proximad of the junction of basal vein and Cu. 

Remarks. This subgenus contains, apart from Apis mellifera, with its race 
A. m. indica Fabr., the large Apis dorsata Fabr. of India and the small A . florea Fabr. 
of India. 

The fossil forms from the Pleistocene belong to the A. mellifera group, and those 
from the Miocene also are rather closer to this Recent species than to either of the 
other two. 


Apis (Apis) armbrusteri Zeuner 1931 

1929 'Scheuthle Biene' Armbruster : 4. 
1931 'Scheuthle Biene' Statz : 44, 46. 
♦1931 Apis armbrusteri Zeuner : 1292, fig. 21, pi. 8, fig. i. 
1938 Hauffapis scheuthlei Armbruster : 43, 92, figs 1-5, 7, 8, 14, 20, 23, 26, 29, 33, 35, 38-40, 
51, 53. 55. 58, 63-65, 67, 70-72, 76, 77, 79. 

HoLOTYPE. Wurttembergische Naturaliensammlung, Stuttgart, Boettingen Col- 
lection No. 47, individual No. 3 (head, thorax and abdomen). Cast : British 
Museum (Natural History) In. 38859 (thorax) and In. 38856 (abdomen). 

Paratypes. Wurttembergische Naturaliensammlung, Stuttgart, Boettingen 
Collection No. 47, individuals Nos i, 2, 4-17. Casts : British Museum (Natural 
History) In.38857 (thorax, No. i), In.38858 (thorax, No. 2), In.38860 (thorax. 
No. 4). 

Other material. The specimens described by Armbruster (1938) from the 
Upper Miocene of the Randecker Maar, all here attributed to Apis armbrusteri as 
a species. Subspecific distinctions have, however, been retained as will be seen 

Distribution. Upper Miocene, Wiirttemberg. 

Diagnosis. A bee resembling Apis mellifera in size and appearance, with wax 
mirrors on the abdomen, the posterior edges of the sternites being apparently 
straight. Wings proportionately slightly broader than in the Recent species, position 
of the junction of the second m-cu with M, and shape of the first m-cu variable. 

Remarks. In 1931, one of us (F. E. Z.) described a fossil swarm of honey bees 
from the thermal limestone of Bottingen on the Swabian Alb, Wiirttemberg, under 
the name Apis armbrusteri. It had perished in peculiar circumstances, having 
fallen into a fissure, on the edge of the crater of an Upper Miocene volcano, from 
which a spring was issuing and which released carbon dioxide into the atmosphere. 
Whilst these specimens afforded many details of the body structure, the wing vena- 
tion could not be studied. 

After the termination of the eruptions, but still in the Upper Miocene, lakes had 
formed in the rather larger crater of the Randecker Maar which is situated in the same 
volcanic area. Shales deposited in these lakes produced an abundant insect fauna, 
again probably because either the water or the atmosphere was temporarily poisoned. 
In 1926 Wilhelm Scheuthle discovered bees here. 

In 1928 he was joined by Professor Armbruster, and their search resulted in the 
recovery of a very large number of specimens which, however, were not scientifically 
described until 1938, under the names of Hauffapis scheuthlei, H. scheeri and H. 
scharmanni Armbruster, with a number of subspecies. These bees provide valuable 
information about venation, being compressed in much the same way as the bees 
from Rott. The variation observed is great, but no clear line can be drawn justifying 
the maintenance of species. It is altogether very unlikely that in an area of 50 miles 
there should have existed simultaneously four hardly distinguishable species of 
honey bees with four more subspecies. On geographical, stratigraphical and mor- 
phological grounds, therefore, the Bottingen and Randeck bees should be regarded 


as one variable species. The specific names given by Armbruster have been allotted 
subspecific rank, as they may be useful in the discussion of the variation, whilst 
Armbruster's subspecies have been synonymized with his species. In essence, this 
appears to represent not only ours, but Professor Armbruster's views, who in 1938 
stated that he considered the Randeck bees identical with Apis armbrusteri Zeuner. 
The new names were perhaps, in part, prompted by his wish to honour some of his 
collaborators and colleagues. 

All specimens so far recovered at Randeck and Bottingen are workers, except for 
a single fragment of a drone from Randeck. The following four subspecies may be 

Apis (Apis) armbrusteri armbrusteri Zeuner 1931 

*i93i Apis armbrusteri Zeuner : 292, fig. 21, pi. 8, fig. i. 
1934 Apis armbrusteri Zeuner ; Statz : 7. 
1938 Apis armbrusteri Zeuner ; Armbruster : 16, 45. 

HoLOTYPE AND Paratypes. See under species, above. 

Distribution. Miocene, 'Bottinger Marmor', Bottingen, Swabian Alb, Wiirttem- 

Diagnosis. Head 2 mm, thorax 5 mm, abdomen 12 mm (distended). Meso- 
thorax strongly vaulted, middle segment only slightly sloping. First abdominal 
segment very short, vertical and concave anteriorly. Posterior edges of abdominal 
sternites not or very little angular at mid-line. 

Description. On a piece of red tufa measuring not more than 45 X35 xio mm, 
17 individuals are preserved in a layer and in a closely packed condition. The insects 
appear as undistorted cavities and have to be studied with the aid of casts made in 
an elastic material. The cavities have been broken in various positions, the 
abdomina (being the largest) being available most frequently. In addition, 5 
thoraces and 3 heads can be studied. 

Head. The eyes appear to have been oval, not kidney-shaped. They are 
separated on the vertex, hence the specimens may be regarded as workers. 

Thorax. Almost spherical, strongly vaulted above. Prothorax short, the 
shoulder-buckles not reaching the tegulae. Shoulder-buckles prominent laterally 
in an angular fashion. Mesopleurae curved, about twice as high as long. Middle 
coxae long and fitted into the thorax, not pointing obliquely backwards, but vertical 
to the longitudinal axis of the insect. Mesonotum very strongly vaulted, more so 
than in A. mellifera, and exceeding the scutellum in height. Scutellum sharply set 
off and forming a prominent crosswise flange. Middle segment sloping at a smaller 
angle than in A. mellifera or even in A. florea. 

Abdomen. Stalk short, first segment concave anteriorly, short, as seen from 
above. The posterior margins of the middle sternites are not angular, but more or 
less straight. After careful investigation, this could not be attributed to distension 


or distortion of the abdomen ; it thus appears to be a sound diagnostic character. 
Wax mirrors present. 

Remarks, The Bottingen bees differ from the three Recent species of Apis in 
the shape of the thorax, whose mesonotum is so strongly vaulted that it exceeds 
the scutellum in height, a feature that may be interpreted as a sign of high speciali- 
zation. Similarly, the shape of the first abdominal segment suggests specialization 
beyond the level of Apis mellifera. In other respects, however, the Bottingen bees 
are more primitive than A. mellifera, especially in the shape of the abdominal 
sternites, and perhaps in the slight slope of the middle segment of the thorax. 

It is much to be regretted that no evidence could be obtained about the collecting 
apparatus or the hind legs. Attempts made to expose the legs by carefully grinding 
away the matrix proved unsuccessful. 

A . armhrusteri was a social bee which in all probability lived much like A . mellifera. 
The wax mirrors suggest that combs were built of this substance as is characteristic 
of the genus Apis. The fact that 17 individuals were contained in a closely packed 
condition in the small piece of rock recovered suggests that a much larger number of 
bees perished and were fossilized, a swarm which would have fallen into the death 
trap of the Bottingen fissure and perished almost instantaneously in its poisoned 

Apis (Apis) armhrusteri scharmanni (Armbruster 1938) 
*i938 Hauffapis scharmanni Armbruster : 44, 113, fig. 78. 

HoLOTYPE. Armbruster's specimen and fig. no. 78, in the collection of Dr Hauff, 

Distribution. Upper Miocene, Randecker Maar, Wiirttemberg. 

Diagnosis. Approx. 9-1 mm long ; form compact ; head massive ; abdomen 
plump ; corbicula (tibia) very broad but short, much reduced at base ; apex of 
'pollen kneader' (counterpart of auricle, on the tibia) strong and angular ; wax 
mirrors elongated and hammer-shaped ; vein first R^ possibly not arched. 

Description. This is a small bee though the respective states of preservation 
have to be taken into consideration in comparing it with A . armhrusteri armhrusteri. 
The edge of the tibia, from pollen kneader to the end of the limb, forms almost a 
straight line, as in ^. a. scheuthlei. Second Rs (third submarginal cell) not narrowed 
unduly ; second m-cu joining base of third submarginal cell more towards the 
middle ; cubital basis (according to Armbruster) as long as the corbicula, but re- 
puted to be shorter than in A. a. scheeri ; first m-cu only a little deflected ; Cuj 
straight. The tergites of the abdomen are very narrow but very long. 

Remarks. This is in several respects a curious bee and of the several forms here 
discussed the one perhaps most deserving of specific rank. Unfortunately the type 
appears to be lost ; Armbruster had to use 'photograms' as a substitute for the 
specimen. Other photographs mentioned by the author appear to be of the same 
specimen, and no paratypes are known to exist. 


Apis (Apis) armhrusteri scheeri (Armbruster 1938) 

*i938 Hauffapis scheeri Armbruster : 43, 92, figs 50, 52, 62, 64, 66, 69, 71, 73. 
1938 Hauffapis scheeri var. gallauni Armbruster : 45, figs 31, 59. 
1938 Hauffapis scheeri var. rahdei Armbruster : 45, fig. 50. 

Lectotype. Schempp Collection, Stuttgart-Weil (Armbruster 1938 : fig. 71), 
herein selected. 

Paralectotypes. Collections of Scheuthle and Schempp. 

Other material. Holotypes of Hauffapis scheeri gallauni Armbruster, No. 59 
and of Hauffapis scheeri rahdei Armbruster, No. 50. 

Distribution. Upper Miocene, Randecker Maar, Wiirttemberg. 

Diagnosis. Smaller, plumper than A. a. scheuthlei, with narrower third sub- 
marginal cell, and relatively short, broad and boldly formed hind tibia and basi- 
tarsus. Legs very reminiscent of those of Recent A. mellifera. 

Description. Short dense hair, partly arranged in lines, occurring on the abdo- 
men. The wax mirrors are long and narrow. The first m-cu is distinctly angular 
in the holotype. In most respects this form resembles A. mellifera greatly, and 
especially in the shape and size of the wing, and in certain details of the venation. 

The length of the body varies ; a giant form was given the name of H. s. gallauni 
and a dwarf form H. s. rahdei. There is also much variation in overall size and 
proportions. Armbruster gave the average length of the cubital basis as 2-96 mm, 
though he illustrated specimens in which this measurement is 3-05, 3-2 and 3-3 mm, 
which thus appears to be of little taxonomic value. 

Remarks. Of this very variable subspecies, 24 specimens have been recorded. 
Of these, three are only doubtfully placed here. The total includes the 'giant' and 
'dwarf forms to which Armbruster gave special names. 

Apis (Apis) armhrusteri scheuthlei (Armbruster 1938) 

1929 'Scheuthle Biene' ; Armbruster : 4. 

193 1 Apis sp. ; Zeuner : 296. 

*i938 Hauffapis scheuthlei Armbruster : 43, figs 63, 65, 67, 70, 72, 76, 77, 79. 

1938 Hauffapis scheuthlei var. seemanni Armbruster : 45, figs 8, 11, 21, 26, 53. 

1938 Hauffapis scheuthlei var. zeuneri Armbruster : 45, figs 36, 38. 

Lectotype. W. Scheuthle Collection (Armbruster 1938 : fig. 63), herein selected. 

Paralectotypes. Armbruster, Scheuthle, Schempp and Hauff Collections. 

Other material. Holotype of Hauffapis scheuthlei seemanni (Armbruster Coll. 
no. 8), and paratypes (nos 11, 21, 26, 53). Holotype of Hauffapis scheuthlei zeuneri 
(Armbruster Coll. no. 36), paratype (same collection, no. 28). 

Distribution. Upper Miocene, Randecker Maar, Wiirttemberg. 

Diagnosis. Larger than A . a. scharmanni and A . a. scheeri, size slightly exceeding 
A. mellifera except in dwarf specimens. Third submarginal cell broader than in 
A. a. scheeri. Legs slender, somewhat resembling those of A. dorsata. 


Description. This is the largest of the three forms of bees from Randecker 
Maar here recognized. Though the majority of the specimens are of the size of 
large Recent A. mellifera, giant and dwarf forms have been described. A. s. var. 
seemanni is a giant form, whilst the smallest specimens were named A. s. var. 
zeuneri. The body is covered in hair which is somewhat coarse. 

Compared with A . a. scheeri, the legs oiA. a. scheuthlei are longer and more slender. 
The hind tibia (corbicula), considered by Armbruster to be slightly curved in outline, 
is less narrow towards the base. The basitarsus, too, has a straighter outline. 
Armbruster is inclined to regard the legs as reminiscent of A. dorsata. 

The first submarginal cell of the forewing often appears arched to meet C ; the 
second and third submarginal cells are broader than in ^. a. scheeri ; pterostigma 
much reduced ; the first recurrent vein is markedly angular, with a small stump of 
a vein at the apex of the angle. 

Remarks. Armbruster distinguished two subspecies, A. s. seemanni and A. s. 
zeuneri. These are here synonymized with A. a. scheuthlei as they are merely 
extreme size variants which would be found in any large population. Regarding 
one of the specimens assigned to ^. s. seemanni (no. 35), Armbruster (1938 : 103-4) 
assigned its counterpart to ^. a. scheuthlei (no. 33) and withdrew the identification 
of no. 35 (1938 : 114), without, however, altering the text on the earlier pages. That 
this should have been possible illustrates how small the difference is between these 
size classes. 

Armbruster attached some importance to the length of the cubital basis, i.e. the 
base vein of cells first R^, first Rs and second Rs. He did so presumably because 
it is a measure that can be taken on a comparatively large number of wings, includ- 
ing those the total length of which cannot be measured. In yl. a. scheuthlei this 
cubital basis is reputed to equal the length of the corbicula. In fact it varies from 
3-06 to 3-40 mm. In his A. s. seemanni, the value is 3-67 mm. Of ^. s. zeuneri no 
measurements are given, except for no. 64 (Scheuthle Coll.) which is regarded as 
possibly referable to this form. Here it is 'apparently hardly longer than 3-0 mm'. 
The unreliability of this measurement has been clearly demonstrated by Alber 
(1949), who studied the asymmetry of the wings of bees. 

Altogether 44 specimens have been assigned to this variable 'species', which 
Armbruster regarded as 'possibly, nay even probably' overlapping with Apis 
armbrusteri Zeuner. The wax mirrors are preserved on many of the Randecker 
Maar specimens, but their shape is not discussed. From the figures it appears that 
the margins of the sternites were more or less straight, and the wax mirrors rect- 
angular, as in the Bottingen form. 

Apis (Apis) tnelisuga (Handlirsch 1907) 

♦1907 (Apidae) melisuga Handlirsch : 893. 

HoLOTYPE. Collection of Count Bosnianski, with counterpart ; whereabouts 


Distribution. Miocene, Gabbro, Italy. 

Diagnosis. i8 mm long. Other features believed to be much as in Apis 

Description. Proboscis at least as long as head. Hind leg with distinct bristles 
along the edges. Venation not identifiable. 

Remarks. The broadened hind tibia and tarsus strongly suggest a true Apis. 
It is impossible to decide, however, how far it resembles the Recent A. dorsata, or 
differs from A. a. scheuthlei. It is, therefore, retained as a separate species, in the 
hope that fresh material will become available in the future. 

Apis (Apis) mellifera Linnaeus 1758 
(PI. 4. fig. 6) 

*i758 Apis mellifera Linnaeus : 343, 575. 
1890 Apis mellifica Linnaeus ; Foord : 94, fig. i. 
1909c Apis mellifera Linnaeus ; Cockerell : 317. 

Specimens studied. British Museum (Natural History), Palaeontology Dept., 
58516, 58576. 

Distribution. Pleistocene, East African copal. 

Description. The specimens here listed from East African copal do not differ 
from Recent A. mellifera in any essential point. Body length of specimen no. 58576 
is II mm ; thorax 3-25 mm long and 4 mm wide ; abdomen 6 mm long and 4 mm 
wide. Length of wings 875 mm. 

Remarks. Several honey bees are known from the copal of East Africa. Foord 
(1890) mentioned two specimens side by side. His poor figure suggests the shape 
of the basitarsus being identical with A . mellifera. 

Cockerell (1909c) examined two specimens from the Cambridge Collection. They 
were at first believed to be enclosed in Yarmouth amber and hence to be of Eocene 
age, but he considered it much more likely that sailors had brought them from East 
Africa for the Yarmouth trade in amber fossils. 

Specimen no. 58576 has been studied in detail. Its wing venation is remarkable 
in two respects ; the first submarginal cross vein (first abscissa of Rs) is S-shaped, 
and vein first m-cu (first recurrent) appears to lack the usual angularity. The 
peculiar shape of the submarginal cross vein is known also in Synapis, as well as in 
A. florea and to a lesser extent in A. dorsata. One might, therefore, regard it as a 
primitive character, but since it occurs also in a specimen of A. mellifera from 
Entebbe, Uganda, it appears to have persisted in some East African bees. The 
lack of angularity of first m-cu is believed to be a mere accident of preservation ; 
angularity indeed probably does exist, though it may not be very marked, and it is 
masked in this specimen by a slight fold in the wing. 

The abdomen of the East African copal bee has yellow bands, like its modern 


Family APIDAE incertae sedis 
'Apis* aquitaniensis de Rilly 1930 

*i93o Apis aquitaniensis de Rilly (not seen). 

1931 Abeille mellifere ; Alphandery : 3, fig. on p. 3. 

1938 Apis aquitaniensis de Rilly ; Armbruster : 88. 

1950 Apis aquitaniensis de Rilly ; de Rilly : 45. 

HoLOTYPE. Museum Marseille, No. 5979, as stated by both Armbruster (1938) 
and Roussy (personal communication). Repeated enquiries, however, have not 
succeeded in eliciting its present whereabouts. 

Distribution. Oligocene (Aquitanian) : Aix-en-Provence. 

Diagnosis. Length of body i6 mm, length of thorax 5 mm, length of abdomen 
9 mm, length of forewing 10 mm, length of hind leg 7-5 mm. 

Description. According to Alphandery, two antenna are visible on the head, 
and one eye is well preserved. The rest of the head is badly crushed but its general 
shape is triangular, like heads of bees with mandibles in general. The thorax is 
also crushed and has a black depression in the centre. On its edges there are two 
orange-coloured ridges. The left wing is beautifully spread and shows all the 
characteristics of a honey bee. The third pair of legs is well spread out, and the 
tibia shows a broadening suggestive of a pollen basket. Five abdominal segments 
can be distinguished. 

This description is an almost literal translation from Alphandery (1931). 

Remarks. This specimen was first mentioned in an article by F. de Rilly (1930) 
entitled L'aheille fossile frangaise, of which nothing further is known. It may be 
identical with an article said to have been written by that author in 1924, which 
we have not seen either. Alphandery 's description is vague. His reference to the 
left wing showing 'all the characteristics of the honey bee' is perplexing when applied 
to an Oligocene species, and his figure is useless. Armbruster, aware of this, was 
fortunate enough to receive a facsimile in coloured plastic from F. de Rilly. This, 
however, was scarcely the likeness of an Apis. 

Through the kindness of Monsieur Roussy we have been able to inspect a similar 
facsimile and we agree with Armbruster's conclusion. But the question arises as 
to whether the fault lies with the facsimile. 

A paper by de Rilly (1950) appears to answer this question. In it he asserts once 
again his claim that the specimen under discussion is an Apis, but with 'quelques 
caracteres archaiques (cellule supplementaire de I'aile, eperon tibial)'. According 
to an enlargement of the wing (published in 1924) and reproduced in facsimile, 
however, the supplementary cell is that formed by a cross vein to the marginal 
cell, which is thus a cell completely unknown in Apidae. Moreover, the pterostigma 
is exceptionally large and deep. The eperon-tibial, which undoubtedly belongs to 
the hind tibia (since the tibial spur of the middle leg, being always present in Apis, 
affords no diagnostic character) is worth something, in that it rules out the genus 
Apis proper. The few positive indications which have been made available in de 


Rilly's work (1950) prove that this specimen is not an Apis, and probably not even 
a member of the Apidae. 

*Apis' dormitans Heyden 1862 

*i862 Apis dormitans Heyden : 76, pi. 10, fig. 8. 

1891 Apis dormitans Heyden ; Scudder : 689 (5927). 

1907 Apis dormitans Heyden ; Cockerell : 228. 

1907 Apis dormitans Heyden ; Handlirsch : 892. 

1908 Apis dormitans Heyden ; Handlirsch : 1357. 
191 5 Apis (?) dormitans Heyden ; Buttel-Reepen : 13. 
191 5 Osmia dormitans (Heyden) ; Buttel-Reepen -.14. 
1 91 5 Apis dormitans Heyden ; Meunier : 210. 

193 1 Osmia dormitans (Heyden) ; Statz : 42. 
193 1 Apis dormitans Heyden ; Statz : 42. 

HoLOTYPE. British Museum (Natural History), 58778. 

Distribution. Oligocene (Aquitanian) : Rott am Siebengebirge. 

Remarks. According to Carl von Heyden three specimens existed but the first 
was very poorly preserved and he was far from happy about placing it in the genus 
Apis. The second and third specimens were too meagrely described for recognition. 
Buttel-Reepen considered that the whole habitus, the rounding off of the abdomen 
and the great breadth of the thorax compared with the breadth of the head, 
indicated, not an Apis, but an Osmia. Meunier considered it to be some kind of 
worker bee, but referred to it as a very enigmatical form. All authors agree that it 
has not been proved to be an Apis. 

The re-examination of the holotype, which was damaged by water during the 
war, confirms the views of previous authors. No character is preserved which would 
justify placing it in Apis, and the rounded end of the abdomen precludes it from being 
included in this genus. The venation, which Heyden also shows in his figure, is too 
poor for generic identification. 

Two further specimens of 'A. dormitans', from Rott, now in the Museum of 
Comparative Zoology, Harvard University, were investigated by Cockerell (1907). 
They are specifically different from the specimens referred to above but are legitimate 
members of Apis {Synapis). Hence they require a new name, and have been 
described as Apis {Synapis) henshawi dormiens n. subsp. (p. 241). 

'Apis' styriaca Pongracz 1931 

*i93i Apis styriaca Pongracz : 105. 
1938 Apis styriaca Pongrdcz ; Armbruster : 88. 

Holotype. Whereabouts not known. 
Distribution. Miocene : Parschlug. 

Remarks. A wing fragment only has been referred to, but not described. Its 
affinities are wholly obscure, and the name must be treated as a nomen nudum. 


*Bofnhus' carbonarius Menge 1856 

1856 Bomhus Latreille ; Menge : 26, 

♦1856 Bomhus carbonarius Menge : 27. 

1886 Bomhus Latreille ; Brischke : 278. 

1891 Bomhus carhonarius Menge ; Scudder : 690 (5950). 

1907 Bomhus carhonarius Menge ; Handlirsch : 891. 

I909d Bomhus carhonarius Menge ; Cockerell : 5. 

1928 Bomhus carhonarius Menge ; Wheeler : 97. 

193 1 Bomhus carhonarius Menge ; Salt : 143. 

Distribution. Eocene : Baltic amber. 

Remarks. Apparently Menge had five specimens in two blocks of amber, but 
only two species were present, which he called B. carhonarius and B. pusillus. No 
diagnosis or description was ever published. 

'Bombus' muscorum Roussy 1937 

non 1935a Bomhus muscorum (Linnaeus) ; Richards : 73. 
♦1937 Bomhus muscorum Roussy : 58. 

Distribution. Eocene : Baltic amber. 
HoLOTYPE. Whereabouts unknown. 

Description. A solitary bee, red and black, and abundantly hairy. No further 

Remarks. Cannot be assigned to any genus on the characters given. 

*Bombus* pusillus Menge 1856 

1856 Bomhus Latreille ; Menge : 26. 
^1856 Bomhus pusillus Menge : 27. 
1891 Bomhus pusillus Menge ; Scudder : 691 (5954). 
1907 Bomhus pusillus Menge ; Handlirsch : 891. 
i909d Bombus pusillus Menge ; Cockerell : 5. 
1928 Bomhus pusillus Menge ; Wheeler : 96. 

Distribution. Eocene : Baltic amber. 
Remarks. See 'Bomhus' carhonarius Menge above. 

'Bombus' antiquus Heyden 1859 

'1859 Bomhus antiquus Heyden : 12, pi. 2, fig. 4. 

1862 Bomhus antiquus Heyden ; Heyden : 75. 

1891 Bomhus antiquus Heyden ; Scudder : 690, pi. 31, fig. 4 (5949). 

1907 Bombus antiquus Heyden ; Handlirsch : 891. 

193 1 Bomhus antiquus Heyden ; Cockerell : 309. 

1936 Bomhus antiquus Heyden ; Statz : 260. 

HoLOTYPE. Whereabouts now unknown. Originally in von Dechen's collection. 


Distribution. Oligocene (Aquitanian) : Rott am Siebengebirge. 

Remarks. Specimen lacking several parts and even Heyden (1862) considered 
it 'perhaps not different from Osmia carbonum', thus making its generic status very 
insecure. There is no useful purpose served in synonymizing it with the latter 
species, since Heyden's statement is far too casual, and possibly had little real 
significance. No subsequent author has considered the specimen to be 'good'. 

*Bombus* grandaevus Heer 1849 

*i849 Bombus grandaevus Heer : 96, pi. 7, figs 3a, b. 

1856 Bombus grandaevus Heer ; Giebel : 182-183. 

1867 Bombus grandaevus Heer ; Heer : 5, pi. 3, figs 6, 7. 

1 891 Bombus grandaevus Heer ; Scudder : 690 (5952). 

1895 Bombus grandaevus Heer ; Scudder : 122. 

1907 Bombus grandaevus Heer ; Handlirsch : 891. 

1928 Bombus grandaevus Heer ; Wheeler : 97. 

193 1 Bombus grandaevus Heer ; Cockerell : 309. 

1938 Bombus grandaevus Heer ; Armbruster : 87. 

HoLOTYPE. Badische Landessammlung fiir Naturkunde, Karlsruhe. Not traced. 
(Heer 1849 : fig. 3.) 

Another specimen. As figured by Heer (1867). 

Distribution. Miocene : Radoboj. 

Remarks. Cockerell regarded this form as generically unclassifiable. Scudder 
thought it was not a Bombus, but possibly belonging to Eucera. Handlirsch (1907) 
also believed that it belonged near Eucera but to a new genus. Heer's illustrations 
provide no help. They suggest, however, that two different specimens were in 
existence, those figured in 1849 ^^^^ 1867. 

We have made enquiries at Karlsruhe and visited the Museum, but the specimens 
cannot be found. It is possible they may come to light when the material packed 
away in the cellars during the war is sorted out. 

' Bombusoides' mengei Motschulsky 1856 

♦1856 Bombusoides Mengei Motschulsky : 28. 
1 89 1 Bombusoides mengei Motschulsky ; Scudder : 691 (5955). 
1907 Bombusoides mengei Motschulsky ; Handlirsch : 892. 
I909d Bombusoides mengei Motschulsky ; Cockerell : 5. 
1928 Bombusoides mengei Motschulsky ; Wheeler : 97. 

Distribution. Eocene : Baltic amber. 

Description. A bumble bee not more than i^ lines (3-175 mm) long. No further 
description provided. 

Remarks. On formal grounds this is the genotype of Bombusoides Motschulsky, 
The holotype is lost. The size given is much too small for any bumble bee. 



Apidae gen. et sp. indet. i (Brischke) 

1886 Bomhus Latreille ; Brischke : 278. 

1 89 1 Bombus Latreille ; Scudder : 690 (5946). 

1907 Bomhus Latreille ; Handlirsch : 891. 

Distribution. Eocene : Baltic amber. 

Remarks. A single 'Bomhus' mentioned but not described. Probably either 
B. carhonarius Menge or B. pusillus Menge, both nomina nuda (p. 252). 

Apidae gen. et sp. indet. 2 (Brischke) 

1886 Melipona Brischke : 278. 

1891 Melipona (vie.) Brischke ; Scudder : 716 (6175). 

1896 Melipona Tosi : 356. 

1907 Melipona (vie.) Brischke ; Handlirsch : 892. 

Distribution. Eocene : Baltic amber. 

Description. 'Venation is reminiscent of the South American genus Melipona 
but differs in the structure of the legs.' 

Remarks. Two specimens appear to have been known to Brischke, but the 
described specimens are now lost. The comparison with Melipona means little in 
view of the existence of the meliponoid Electrapis. 

Apidae gen. et spec, indet. 3 (Burmeister) 

1832 Trigona Burmeister : 636. 

1836 Trigona Burmeister : 577. 

1856 Trigona Burmeister ; Giebel : 184. 

1891 Trigona (?) Burmeister ; Scudder : 733 (6324). 

1896 Trigona Tosi : 356. 

1907 Trigona (?) Burmeister ; Handlirsch : 892. 

Distribution. Eocene : Baltic amber. 

Remarks. Perhaps identical with Electrapis [Roussyana] palmnickenensis 
(p. 233). Specimen lost. 

Apidae gen. et sp. indet. 4 (Scudder) 


Bomhus Scudder : 



Bomhus Scudder : 

: 280. 


Bomhus Scudder : 

: 690 (6947)- 


Bomhus Scudder ; 

; Handlirsch 

Distribution. Oligocene : Florissant. 

Remarks. One of the specimens of bees which Scudder knew from Florissant 
was assigned to Bomhus without further description or comment. 



Lithoblatta lithophila (Germar 1839) 

839 Musca lithophila Germar : 222. 

864 Blattidium beroldingianum Heer : 300, fig. 8. 

869 Musca lithophila Germar ; Weyenbergh : 256, pi. 34, fig. 2 (no. 6511). 

869 Bombus (?) conservatus Weyenbergh ; Weyenbergh : 259, pi. 34, fig. 7. 

869 Apiaria veterana Weyenbergh ; Weyenbergh : 260, pi. 34, fig. 8 (no. 6480 nee 6431). 

886 Mesoblattina lithophila (Germar) ; Scudder : 464. 

886 Mesoblattina lithophila (Germar) ; Deichmiiller : 6, pi. i, figs 1-6. 

888 Mesoblattina lithophila (Germar) ; Oppenheim : 221, pi. 30, figs 6, 8. 

895 Mesoblattina lithophila (Germar) ; Meunier : 223. 

898 Mesoblattina lithophila (Germar) ; Meunier : pi. 25, figs 76-78 ; pi. 26, figs 85-86 ; 

pi. 27, figs 92-93. 

898 Naucoris carinata Meunier : pi. 27, fig. 88. 

907 Lithoblatta lithophila (Germar) ; Handlirsch : 530, pi. 46, fig. 7. 

931 Bombus conservatus (Weyenbergh) ; Cockerell : 309. 

HoLOTYPE. Whereabouts not known, probably lost. 

Specimens studied. Teyler Museum, Haarlem, Netherlands, No. 15324 {Bombus 
conservatus) and No. 15305 {Apiaria veterana). 

Distribution. Upper Jurassic : Solnhofen, Bavaria. 

Remarks. Meunier regarded both B. conservatus and A. veterana as cockroaches 
{Mesoblattina). His view was shared by Handlirsch who identified them with 
Germar 's Musca lithophila and erected the new genus Lithoblatta. Thus the sup- 
posed bees of the Upper Jurassic have proved to be Orthoptera. 

The specimens in Haarlem which we have studied support this view. No. 15324 
has an orthopteroid habitus, and No. 15305 has long antennae and orthopteroid 
tibia. Cockerell, who saw B. conservatus, also held that it is 'wholly obscure and 
there is no reason for regarding it as a bee'. 

VI. appendix 
by S. F. Morris 

CoUetidae gen. et sp. indet. 
1927 Colletidae ; Blair : 139. 
Material. Whereabouts unknown. 

Distribution. According to Blair the material is 'perhaps Upper Pliocene, 
perhaps Quaternary or probably Sub-recent'. Oil sands ; near Fyzabad, Trinidad. 

Ctenoplecfrella splendens Kelner-Pillault 1970 
*i97oa Ctenoplectrella spendens Kelner-Pillault : 13, pi. i, figs 3, 4. 

Material. Holotype : Geologisch-palaeontologischen Institut der Universitat, 
Gottingen. Paratype : No. 49, Zoologisches Museum der Humboldt-Universitat, 

Distribution. Eocene, Baltic amber. 


Anthidiini gen. et sp. indet. 
1972 Anthidiini ; Sando : 424. 
Material. Gerard Germs Collection, 

Distribution. Recent (collected from rocks of Precambrian and Lower 
Palaeozoic age). 

Osmia sp. 

1972 Osmia sp. ; Sando : 421. 

Material. United States National Museum ; United States Geological Survey, 
Glacier National Park Museum ; Harvard University, Paleobotany Collection. 

Distribution. Recent (in rocks of Precambrian to Upper Palaeozoic age), 
Montana, Wyoming. 

Apoidea (? Anthophorinae) sp. 

1969 Apoidea (? Anthophorinae) sp. ; Werner : 12, pi. 19, fig. 3. 
Material. United States National Museum 257, B.218. 
Distribution. Early Pleistocene, Rita Blanca lake deposits. 

Probombus hirsutus Piton (ms) 

1940 Probombus hirsutus Piton (ms) : 218. 

1969b Probombus hirsutus Piton ; Kelner-Pillault : 526 {nomen nudum). 

Distribution. Eocene, Menat, Puy-de-D6me, France. 

Trigona (Hypotrigona) eocenica Kelner-Pillault 1970 
♦1970b Trigona {Hypotrigona) eocenica Kelner-Pillault : 437, figs 1-3. 

Material. Holotype : Geologisch-palaeontologischen Institut der Universitat, 
Gottingen. Paratype formerly in the Geological Museum of the University, 
Konigsburg ; now in the Humboldt Museum, Berlin. 

Distribution. Eocene. Baltic amber. 

Trigona (Hypotrigona) dominicana Wille & Chandler 1964 

♦1964 Trigona [Liotrigona) dominicana Wille & Chandler : 188, figs 1-5. 

Material. Holotype and five paratypes in the collection of the Illinois Natural 
History Survey. 

Distribution. Oligocene, Dominican Republic. 


Trigona (Nogueirapis) silacea Wille 1959 

*i959 Trigona {Nogueirapis) silacea Wille : 849, pi. 119, text-figs 1-4. 
1964 Trigona {Nogueirapis) silacea Wille ; Wille : 120, fig. i. 

Material. Holotype : 12601. Paratypes : 12602-11. In University of Cali- 
fornia Museum. 

Distribution. Middle Miocene, Simjovel, Chiapas Province, Mexico. 

Electrapis minuta Kelner-Pillault 1970 
♦1970a Electrapis minuta Kelner-Pillault : 16, pi. 2, figs 3, 5B. 

Material. Syntypes : Geologisch-palaeontologischen Institut der Universitat, 
Gottingen ; Block 50a, Zoologisches Museum der Humboldt-Universitat, Berlin. 
Distribution. Eocene, Baltic amber. 

Apis catanensis Roussy i960 

♦1960 Apis catanensis avolii Roussy : 8, fig. 2. 
1969b Apis catanensis avolii Roussy ; Kelner-Pillault : 524. 

Material. One specimen in the private collection of Anastasi and Guiseppe 
Avolio, jewellers of Sicily. 

Distribution. Miocene, near Catania, Sicily. 

Remarks. Roussy used the trinomen Apis catanensis avolii, but since the species 
was not split the third name is superfluous. 

? Apoidea sp. 

1 82 1 Vast numbers of bees and nests ; Henslow:50i. 
Material. Whereabouts unknown. 
Distribution. Parish of Kirk Balaft, Isle of Man ; ? Pleistocene. 

Not Apoidea 

1962 Native bee ; Carroll : 264, fig. i. 
Material. Melbourne University, Geological Dept., M.U.G.D. 3518. 
Distribution. ? Lower Jurassic, Koonwarra, South Gippsland, Victoria, 

VII. acknowledgements 

The task of an amanuensis is a difficult one, but to undertake the publication of a 
manuscript on a subject with which one is unfamiliar, and whose authors one did 
not work with or even know, requires a particular courage and dedication. Thus, 


great credit is due to Mr S. F. Morris for his initiative in revivifying the manuscript 
and in undertaking much of the editorial work towards its pubHcation. Thanks 
are also due to Mr D. L. F. Sealy who prepared the paper for the press, and to 
Professor O. W, Richards for his advice. 

H. W. Ball 

Keeper of Palaeontology 


[Although every effort has been made to trace all the references cited in the text, in a few cases, indi- 
cated below by *, this has unfortunately not been possible; though bibliographic references can be given 
in most instances, they must be cited as 'not seen' (by the Editor) . It is evident that some works which 
were available to the authors are not now to be obtained in any British library. The Editor gratefully 
acknowledges the assistance of the librarians of the British Museum (Natural History) in his efforts to 
trace the missing works. Though not cited in the text, references marked f are known to have been 
consulted by the authors during the preparation of this paper. They are included as a contribution to 
the general bibliography of the subject. S. F. M.] 

Alber, M. 1949. Asymmetrie der Bienenfiiigel. Arch. Bienenk., Berlin, 26 : 14-16. 
*Alphandery, E. 1931. Traite complet d' apiculture. vi + 572pp. Paris. 

Armbruster, L. 1929. Wilhelm Scheuthle f . Bienenpflege, Wiirttemberg, 51 (i) : 5-8. 
* 1938. Versteinerte Honigbienen aus dem obermiocanen Randecker Maar. Arch. 

Bienenk., Berlin, 19 : 1-48, 97-133, figs 1-79. 
AsHMEAD, W. H. 1899. Classification of the Bees, or the Superfamily Apoidea. Trans. Am. 

ent. Soc, Philadelphia, 26 : 49-100. 
Bequaert, J. C. & Carpenter, F. M. 1941. The antiquity of social insects. Psyche, 

Camh. 48 : 50-55. 
Berthold, A. A. 1827. Latreille's naturliche Familien des Thierreichs. viii + 602 pp. 

fBiscHOFF, H. 1927. Biologic der Hymenopteren. 598 pp. Berlin. 
Blair, K. G. 1927. Insect Remains from Oil Sand in Trinidad. Trans, ent. Soc. London 

75: 137-141. 
Brischke, D. 1886. Die Hymenopteren des Bernsteins. Schr. naturf. Ges. Danzig (n.f.) 

6 (3) : 278-279. 
Brown, R. W. 1934. Celliforma spirifer ; the fossil larval chambers of mining bees. /. 

Wash. Acad. Sci., Menasha, 24 : 532-539, 5 figs. 
1935- Further notes on fossil larval chambers of mining bees. /. Wash. Acad. Sci., 

Menasha, 25 : 526-528. 
1941a. The comb of a wasp nest from the Upper Cretaceous of Utah. Am. J. Sci., 

New Haven, 239 : 54-56, i pi. 

1941b. Concerning the antiquity of social insects. Psyche, Camh. 48 : 105 -no. 

Burmeister, C. H. C. 1832. Handbuch der Entomologie 1. xvi + 698 pp., 16 pis. Berlin. 

1836. A manual of Entomology, xii + 654 pp., 33 pis. London. 

fBuTTEL-REEPEN, H. VON 1903. Die Stammesgeschichtliche Entstehung des Bienenstaates 

sowie Beitrdge zur Lebensweise der solitdren und sozialen Bienen. 138 pp., 20 figs. Leipzig. 
1906. Apisticae. Beitrage zur Systematik, Biologic, sowie zur geschichtlichen und geo- 

graphischen Verbreitung der Honigbiene {Apis mellifica L.), ihrer Varietaten und der 

librigen Apis-Arten. Mitt. zool. Mus. Berl. 3 : 1 17-201, 8 figs. 

1915. Leben und Wesen der Bienen. 300 pp. Brunswick. 

Buxton, P. A. 1932. Ancient workings of insects, perhaps bees, from Megiddo, Palestine. 

Proc. ent. Soc. Lond. 7 (i) : 2-4. 
Carroll, E. J. 1962. Mesozoic fossil insects from Koonwarra, South Gippsland, Victoria. 

Aust. J. Sci., Sydney, 25 : 264-265, 2 figs. 


CocKERELL, T. D. A. 1900. Observations on bees collected at Las Vegas, New Mexico, and 
in the adjacent mountains. Ann. Mag. nat. Hist., London, {7) 5 : 401-416. 

1906. Fossil Hymenoptera from Florissant, Colorado. Bull. Mus. comp. Zool. Harv. 

50 : 33-58. 

1907. A Fossil Honey-Bee. Entomologist, London, 40 : 227-229. 

1908a. Descriptions and Records of Bees, 19. Ann. Mag. nat. Hist., London, (8) 1 : 


1908b. Descriptions and Records of Bees, 20. Ann. Mag. nat. Hist., London, (8) 2 : 


1908c. A fossil leaf-cutting bee. Can. Ent., Ottawa, 40 : 31-32. 

i9o8d. Some results of the Florissant expedition of 1908. Am. Nat., Salem, Mass., 

42 : 569-581, 12 figs. 

1909a. Two fossil bees. Ent. News, Philadelphia, 20 : 159- 161. 

1909b. A Catalogue of the Generic Names based on American Insects and Arachnids 

from the Tertiary Rocks with Indications of the Type Species. Bull. Am. Mus. nat. Hist., 
New York, 26 : 77-86. 

1909c. Some European Fossil Bees. Entomologist, London, 42 : 313-317. 

i909d. Descriptions of Hymenoptera from Baltic Amber. Schr. phys.-okon. Ges. 

Konigsh. 50 : 1-20, 14 figs. 

19096. New North American Bees. Can. Ent., Ottawa, 41 : 393-395. 

I909f. Some Additional Bees from Prussian Amber. Schr. phys.-okon. Ges. Konigsh. 

50 : 21-25. 

1911a. Fossil insects from Florissant, Colorado. Bull. Am. Mus. nat. Hist., New York, 

30: 71-82. 

1911b. Descriptions and Records of Bees, 34. Ann. Mag. nat. Hist., London, (8) 

7: 225-237. 

1913a. Some fossil insects from Florissant, Colorado. Proc. U.S. natn. Mus., Washing- 
ton, 44: 341-346, pi. 56. 

1913b. Some fossil insects from Florissant. Can. Ent., Ottawa, 45 : 229-233. 

1914. Miocene fossil insects. Proc. Acad. nat. Sci. Philad. 66 : 634-648, 5 figs. 

1917. New Tertiary insects. Proc. U.S. natn. Mus., Washington, 52 : 373-384, pi. 31. 

1921. Fossil Arthropods in the British Museum. Ann. Mag. nat. Hist., London, (9) 

8: 541-545. 5 figs. 

1922. Fossils in Burmese Amber. Nature, Lond. 109 : 713-714. 

■ 1923. Two Fossil Hymenoptera from Florissant (Vespidae, Megachilidae) . Ent. News, 

Philadelphia, 34 : 270-271. 

1925a. Descriptions and Records of Bees, 106. Ann. Mag. nat. Hist., London, (9) 

16 : 416-423. 

1925b. Tertiary insects from Kudia River, Maritime Province, Siberia. Proc. U.S. 

natn. Mus., Washington, 68 (5) : 1-16, pis 1-2. 
193 1. Insects from the Miocene (Latah) of Washington, II. Hymenoptera and Hemip- 

tera. Ann. ent. Soc. Am., Washington, D.C., 24 : 309-312, pi. i, figs 2, 4, 8. 
1934. Some African Meliponine Bees. Revue Zool. Bot. afr., Brussels, 26 : 46-62, 

9 figs. 
Curtis, J. 1828. British Entomology 5 : pis 195-241 with explanations. London. 
Dall, W. H. 1915. A monograph of the molluscan fauna of the Orthaulax pugnax zone of 

the Oligocene of Tampa, Florida. Bull. U.S. natn. Mus., Washington, D.C., 90: 1-173, 

pis 1-26. 
fDALLA Torre, K. W. von & Friese, H. 1899. Die hermaphroditen und gynandromorphen 

Hymenopteren. Ber. naturw.-med. Ver. Innsbruck 24 : 96 pp., i pi. 
Daly, H. V. 1973. Bees of the Genus Ceretina in America North of Mexico (Hymenoptera : 

Apoidea). Univ. Calif. Pubis Ent., Berkeley, 74 : 1-114, 2 pis. 
Deichmuller, J. V. 1886. Die Insecten aus dem lithographischen Schiefer im Dresdener 

Museum. Mitt. K. miner. -geol. prdhist. Mus. Dresden 7 : 1-88, pis 1-5. 


DouRS, R. Z. 1873. Hymenopteres du Bassin Mediterraneen. Revue Mag. Zool., Paris, 

(3) 1 : 274-325. 
Fabricius, J. C. 1775. Sy sterna entomologiae. 30 + 832 pp. Flensburg and Leipzig. 

1805. Sy sterna Piezatorum. 439 + 30 pp. Brunswick. 

FooRD, A. S. 1890. Notes on a collection of East Coast amber belonging to Mrs Burwood of 

Yarmouth. Trans. Norfolk Norwich Nat. Soc. 5 : 92-95, i pi. 
•fFRiESE, H. 1923. Die europaischen Bienen [Apidae). 456 pp., 33 pis. Berlin and Leipzig. 
Germar, E. F. 1839. Die versteinerten Insecten Solenhofens. Nova Acta physico-med., 

Breslau and Bonn, 19 (i) : 187-222, pis 21-23. 
1849. Ueber einige Insekten aus Tertiarbildungen. Z. dt. geol. Ges., Berlin, 1 : 52-66, 

pi. 2. 
GiEBEL, C. G. 1856. Fauna der Vorwelt mit stater Beriicksichtigung der lebenden Thiere. 

511 pp. Leipzig. 
Handlirsch, a. 1906-8. Die fossilen Insekten und die Phylogenie der rezenten Formen. 

1430 pp., 51 pis. 

f 1921. In Schroder, C., Handbuch der Entomologie 3 (7) : 117-304, figs 52-327. Jena. 

Heer, O. 1849. Die Insektenfauna der Tertiargebilde von Oeningen und von Radoboj. 

Neue Denkschr. Allg. schweiz. Ges. ges. Naturw., Zurich, 11 : 1-264. 
f 1850. Die Insektenfauna der Tertiargebilde von Oeningen und von Teningen und von 

Radoboj in Croatien, II. Neue Denkschr. Allg. schweiz. Ges. ges. Naturw., Zurich, 11 : 

264 pp., 17 pis. 

1864. Ueber die fossilen Kakerlaken. Vjschr. naturf. Ges. Ziirich 9 : 273-302, i pi. 

1865. Die Urwelt der Schweiz. 622 pp., 11 pis, 368 figs. Zurich. 

1867. Fossile Hymenopteren aus Oeningen und Radoboj. Neue Denkschr. Allg. 

schweiz. Ges. ges. Naturw., Zurich, 22 : 1-42, 3 pis. 

1872. Le Monde primitif de la Suisse. 801 pp., 11 pis, 368 figs. Geneva and Basle. 

1876. The Primaeval World of Switzerland, 2. 324 pp., 11 pis, 370 figs. London. 

1879. Die Urwelt der Schweiz. 2nd (rewritten and enlarged) ed. xix + 713 pp., 12 pis, 

417 figs. Zurich. 
Henslow, J. S. 1 82 1. Supplementary observations to Dr. Berger's account of the Isle of 

Man. Trans, geol. Soc. Lond. 5 : 482-505. 
Heyden, C. von 1859. Fossile Insekten aus der Rheinischen Braunkohle. Palaeonto- 

graphica, Cassel, 8 : 1-15, pis 1-2, figs 1-13. 
1862. Gliedertiere aus der Braunkohle des Niederrhein's, der Wetterau und der Rohn. 

Palaeontographica, Cassel, 10 : 62-82, pi. 10. 
HuNEFELD, L. 1831. Derselbe iiber Bernsteininsecten. Isis, Leipzig 15: col. iioo (numb. 

in error 2000). [Determinations by C. H. C. Burmeister.] 
f Jack, R. W. 1917. Parthenogenesis amongst the Workers of the Cape Honey Bee : Mr 

G. W. Onions' Experiments. Trans, ent. Soc. London 49 : 396-403, pis 105-106. 
f Jones, F. W. 1943. Habit and Heritage. 100 pp. London. 
JuRiNE, L. 1807. Nouvelle mdthode de classer les Hymenopteres et les Dipteres 1. 319 pp., 

14 pis. Geneva. 
fKELNER-PiLLAULT, S. 1968. Lcs Abcilles fossiles. 13th Int. Congr. Ent., Proc, Moscow, 

1 : 256. 
1969a. Abeilles fossiles ancetres des Apides sociaux. Proc. VI Congr. lUSSI, Berne : 


1969b. Les abeilles fossiles. Memorie Soc. ent. ital., Genoa, 48 : 519-534, 10 figs. 

1970a. L'ambre balte et sa faune entomologique avec description de deux Apoides 

nouveaux. Annls Soc. ent. Fr., Paris (n.s.) 6 : 3-24, 6 figs. 

1970b. Une Melipone (s.l.) de I'Ambre balte (Hym. Apidae). Annls Soc. ent. Fr., Paris, 

(n.s.) 6 : 437-441, 3 figs. 

Kerr, W. E. & Maule, V. 1964. Geographical distribution of stingless bees and its impli- 
cations (Hymenoptera : Apidae). Jl N.Y. ent. Soc, New York, 72 : 2-8, 4 figs, 6 maps. 

KiRBY, W. 1802. Monographia Apum Angliae 1. xxii + 258 pp., 14 pis. Ipswich. 


Kohl, F. F. 1894. Zur Hymenopterenfauna Afrikas. Annlnnaturh. Mus.Wien9 : 2'jg-^^o, 

pis 13-17. 
Latreille, p. a. 1802a. Histoire naturelle des Fourmis, et recueil de mimoires et d' observations 

sur les Aheilles, les Araignees, les Fauchers, et autres Insectes. xvi4-445 pp., 12 pis. Paris, 
f 1802b. Histoire naturelle, generate et particuliere des Crustaces et des Insectes 3. 468 pp. 


1803. Nouveau Dictionnaire d'Histoire Naturelle 18. 595 pp., pis 23-33. Paris. 

1804. Nouveau Dictionnaire d'Histoire Naturelle 24. Tableaux Methodiques .... 

238 pp. Paris. 

1809. Genera Crustaceorum et Insectorum 4. 399 pp. Paris. 

1810. Considerations generates sur I'ordre naturel des Animaux composant les classes des 

Crustaces, des Arachnides, et des Insectes ; avec un Tableau methodique de leurs genres, 

deposes en families . 444 pp. Paris. 
1825. Families naturelles du Regne Animal, exposees succinctement et dans un ordre 

analytique avec Vindication de leurs genres. 570 pp. Paris. 
Lea, a. M. 1925. Notes on some calcareous insect puparia. Rec. S. Aust. Mus., Adelaide, 

3 : 35-36, I pi. 

Lepeletier de Saint-Fargeau, a. L. M. 1841. Histoire naturelle des Insectes. Hymenop- 

teres, 2. 680 pp. Paris. 
& Audinet-Serville, J. G. 1828. In Latreille, P. A., Encyclopedic methodique. 

Histoire naturelle 10. 833 pp. Paris. 
fLEUENBERGER, F. 1931. Fine fossile Biene. Schweiz. Bienenztg, Aarau, 3 : 3 pp., i fig. 
Linnaeus, C. 1758. Systema naturae .... loth edition, 1. 824 pp. Holmiae. 
Magretti, p. 1884. Risultati di raccolte imenotterologische nell' Africa orientale. Annali 

Mus. civ. Stor. nat. Giacomo Doria, Genoa, 21 : 523-636, pi. i. 
1895. Imenotteri. Annali Mus. civ. Stor. nat. Giacomo Doria, Genoa, 35 : 149-173, 

4 figs. 

Malfatti, G. i88ia. Bibliografia degli Insetti fossili Italiani finora conosciuti. Atti Soc. 

ital. Sci. nat., Milan, 24 : 89-100. 
f- i88ib. Due piccoli imenotteri fossili dell'Ambra siciliana. Atti Accad. Lincei Transunti, 

Rome, (3) 5 : 80-83, 2 figs, 
f Manning, F. J. 1952. Recent and fossil honey bees : some aspects of their cytology, 

phylogeny and evolution. Proc. Linn. Soc. Lond. 163 : 3-8, i pi. 
i960. A new fossil bee from Baltic amber, nth Int. Congr. Ent., Verh., Vienna, 1 : 306- 

308, pi. 5, fig. I. 
Menge, a. 1856. Lebenszeichen vorweltlicher, im Bernstein eingeschlossener Thiere. 

Programm . . . offentlichen Priifung der Schiller der Petrischule : 1-32. Danzig. 
Meunier, F. 1895. Note sur des empreintes d'lnsectes des schistes de Solenhofen. Bull. 

Soc. ent. Fr., Paris, 64 : ccxxiii-ccxxiv. 
1898. Les insectes des temps secondaires. Archs Mus. Teyler, Haarlem, (2) 6 : 89-149, 

pis 1-30. 
1914. Nouvelles recherches sur quelques Insectes du Sannoisien d'Aix-en-Provence. 

Bull. Soc. gSol. Fr., Paris, (4) 14 : 187-198, pis 6-7, text-figs i-ii. 
1915. tJber einige fossile Insekten aus den Braunkohlenschichten (Aquitanien) von 

Rott (Siebengebirge) . Z. dt. geol. Ges., Berlin, 67 : 205-217, pis 21-25, 6 figs. 
1920. Quelques insectes de 1' Aquitanien de Rott, Sept-Monts (Prusse rhenane). Proc. 

Sect. Sci. K. ned. Akad. Wet., Amsterdam, 22 : 727-737, 2 pis, 8 figs. 
Michener, C. D. 1944. Comparative external morphology, phylogeny and a classification 

of the bees (Hymenoptera) . Bull. Am. Mus. nat. Hist., New York, 82 : 151-326, 246 figs. 
1965. A classification of the bees of the Australian and South Pacific regions. Bull. Am. 

Mus. nat. Hist., New York, 130 : 1-362, 789 figs, 15 pis. 
MoTSCHULSKY, V. 1856. Lcttres . . . a M. Menetries, 3. £itud. ent., Helsinki, 5 : 3-38, i pi. 
Moure, J. S. 1961. A Preliminary Supra-specific Classification of the Old World Meliponine 

Bees (Hymenoptera, Apoidea). Studia ent., Rio de Janeiro, 4 : 181-242. 


Novak, O. 1877. Fauna der Cyprisschiefer des Egerer Tertiarbeckens. Sber. Akad. Wiss. 

Wien 76 : 71-96, pis 1-3. 
Oppenheim, p. 1888. Die Insectwelt des lithographischen Schiefers in Bayern. Palaeonto- 

graphica, Cassel, 34 : 215-247, pis 30-31. 
OusTALET, M. E. 1870. Recherches sur les insectes fossiles des terrains tertiaires de la 

France. Annls Sci. giol. Paris 2 (3) : 1-178, pis 1-6. 
Panzer, G. W. F. 1806. Kritische Revision der Insektenfauna Deutschlands 2. Entomo- 

logischer Versuch die Jiirineschen Gattungen der Linneschen Hymenoptern nach dem Fab- 

riciusschen System zu priifen. 271 pp., 2 pis. Nuremburg. 
PiTON, L. 1940. Paleontologie du gisement iocene de Menat. These, Clermont-Ferrand. 
fPLATH, O. E. 1934. Bumblebees and their Ways. 201 pp., 10 pis. New York. 
PoNGRACZ, A. 1928. Die fossilen Insekten von Ungarn. Annls hist.-nat. Mus. natn. hung., 

Budapest, 25 : 91-194, 55 figs. 
193 1 . Bemerkungen iiber die Insektenfauna von Oeningen nebst Revision der Heer'schen 

Typen. Verh. naturh.-med. Ver. Heidelb. 17 : 104-125, pis 3-11. 
•j-RiCHARDS, O. W. 1935a. Bombus muscorum (Linnaeus) and B. smithianus White (Hym.). 

Trans. Soc. Br. Ent., Southampton, 2 : 73-85, pi. 5. 
1935b. Notes on the nomenclature of the Aculeate Hymenoptera, with special reference 

to British genera and species. Trans. R. ent. Soc. Lond. 83 : 143-176. 
*RiLLY, F. de 1930. L'abeille fossile fran9aise. Apiculteur. [Not seen by authors.] 

* 1950. [title ?] Ruchers (10) : 45. 

RoDECK, H. G. 1938. Type specimens of fossils in the University of Colorado Museum. 

Univ. Colo. Stud. gen. Ser., Boulder, 25 : 281-304. 
RosELLi, F. L. 1939. Apuntes de geologia y paleontologia Uruguayas y sobre insectos del 

Cretaceo del Uruguay o descubrimentos de admirables instintos constructivos de esa epoca. 

Boln Soc. Amig. Cienc. nat. Kraglievich-Fontana, Nueva Palmira, 1 (2) : 32-102, 32 figs. 
Ross, H. H. 1936. The ancestry and wing venation of the Hymenoptera. Ann. ent. Am., 

Washington, D.C., 29 : 99-1 11, 2 pis. 
*RoussY, L. 1937. Flore et Faune de I'ambre fossile de la Mer Baltique. Gaz. apic, Mont- 

favet, 388 : incl. p. 66 & fig. 13 (March 1937). 
i960. Insectes et abeilles fossiles de I'ambre de Sicile. Gaz. apic, Montfavet, 635 : 5-8, 

2 figs. 
Salt, G. 1931. Three Bees from Baltic Amber. Bernstein-Forsch., Berlin and Leipzig, 

2 : 136-147. 
Sandhouse, G. A. 1943. The type species of the genera and subgenera of bees. Proc. U.S. 

natn. Mus., Washington, 92 : 519-668. 
Sando, W. J. 1972. Bee-nest pseudofossils from Montana, Wyoming and South- West 

Africa. /. Paleont., Tulsa, 46 : 421-425, fig. i. 
Schletterer, a. 1 89 1. Hymenoptera in expeditione sub auspicio regii imperii Belgici 

perfecta in regione Africae ad Congo flumen inferius collecta, determinata sive descripta 

ab. Annls Soc. ent. Belg., Brussels, 35 : 1-36, pis 1-2. 
ScHMiEDEKNECHT, O. 1930. Die Hymenoptcren Nord- und Mitteleuropas. x+1062 pp., 

127 figs. Jena. 
Schroder, C. 1913-25. Handbuch der Entomologie 3. viii-f- 1201 pp., 1040 text-figs. Jena. 

[See Handlirsch 1921.] 
fScHULZ, W. A. 1907. Alte Hymenoptera. Berl. ent. Z. 51 : 303-333. 
Schutze, E. 1907. In Branca, W. & Fraas, E., Die Lagerungsverhaltnisse Bunter Breccie 

an der Bahnlinie Donauworth-Treuchtlingen und ihre Bedeutung fiir das Riesproblem. 

Phys. Math. Abh. K. Akad. Wiss. Berlin 2 : 25-26, i pi., figs 22-23. 
ScHWARZ, H. F. 1948. Stingless bees (Meliponidae) of the Western Hemisphere. Bull. Am. 

Mus. nat. Hist., New York, 90 : 1-546, pis 1-8, 87 figs. 
ScuDDER, S. H. 1 88 1. The tertiary lake basin at Florissant, Colorado, between South and 

Hayden Parks. Bull. U.S. geol. geogr. Surv. Territ., Washington, 6 : 279-300. 


ScuDDER, S. H. 1883. The tertiary lake basin at Florissant, Colo., between South and 
Hayden Parks. Rep. U.S. geol. geogr. Surv. Territ., Washington, 12 : 271-293. 

1885. Myriopoda, Arachnoidea und Insecta. In Zittel, K. A., Handhuch der Palaeonto- 

logie : 723-893, figs 894-1109. Munich and Leipzig. 

1886. A Review of Mesozoic Cockroaches. Mem. Boston Soc. nat. Hist. 3 : 439-485, 

pis 46-48. 

1887. Partie i, Mollusca et Arthropoda. In Zittel, K. A., Traite de Paleontologie 2. 

897 pp., 1 126 figs. Paris, Munich and Leipzig, 
f 1890. A classed and annotated bibliography of fossil insects. Bull. U.S. Geol. Surv., 

Washington, 69 : loi pp. 
1 891. Index to the known fossil insects of the world including myriapods and arachnids. 

Bull. U.S. Geol. Surv., Washington, 71 : 744 pp. 

1895. The Miocene Insect-fauna of Oeningen, Baden. Geol. Mag., London, (4) 2 : 116- 

122, pi. 6. 
Smith, F. 1854. Catalogue of Hymenopterous Insects in the . . . British Museum 2 : 199-465. 

1858. Catalogue of the Hymenopterous Insects. /. Proc. Linn. Soc, London, 2 : 42-130, 

pis I, 2. 
Snodgrass, R. E. 1935. Principles of Insect Morphology, ix + 667 pp., 319 figs. London. 
Spinola, M. 1808. Insectorum Liguriae species novae aut rariores, . . . [&c.] 2. ii + 262 + 

V pp., 5 pis. Genoa. 
1838. Compte-rendu des Hymenopteres, recueillis par M. Fischer pendant son voyage 

en Egypte. Annls Soc. ent. Fr., Paris, 7 : 437-546. 
Stadelmann, H. 1895. Beitrage zur Kenntniss der Gattung Melipona sens. lat. 

Sber. preuss. Akad. Wiss., Berlin, Jahrg. 1895 : 615-623, 3 figs. 
Statz, G. 1931. Eine neue Bienenart aus Rott am Siebengebirge. Wiss. Mitt. Ver. Natur-u. 

Heimatk. Koln 1 : 39-60. 
1934- Neue Beobachtungen iiber fossile Bienen aus dem Tertiar von Rott am Siebenge- 
birge. Arch. Bienenk., Berlin, 15 (i) : i-io, 10 figs. 
1936. Ueber alte und neue fossile Hymenopterenfunde aus den tertiaren Ablagerungen 

von Rott am Siebengebirge. Decheniana, Bonn, 93 : 256-312, pis 1-13. 
* 1943. Bienen, Blatter und Bliiten dem rheinischenBraunkohlenwald. Rhein. Heimatphl., 

Diisseldorf, 13 (for 1941) : 100- 112, 14 figs. Also (same title) Dtsch. Imkerfiihrer 16 

(for 1942) : 6 pp., 22 figs. [These works, not seen by the authors, are desiderata in that they 

may contain new taxonomic names.] 

* 1944- [title ?] Umschau, Frankfurt-a.-M., 48 (4) : 63, figs 2, 5. 

Theobald, N. 1937. -^^•^ insectes fossiles des terrains oligocenes de France. 473 pp., 29 pis. 

Timon-David, J. 1943. Premiere etude des Insectes fossiles ohgocenes du basin de Mar- 
seille. C. r. hebd. Seanc. Acad. Sci., Paris, 216 : 455-457. 
- — - 1944- Insectes fossiles de I'OHgocene inferieur des Camoins (Bassin de Marseille) II. 

Hymenopteres. Bull. Soc. ent. Fr., Paris, 49 : 40-45, 5 figs. 
Tosi, A. 1896. Di un nuovo genere di Apiaria fossile nell'ambra di Siciha (Meliponorytes 

succini-M. sicula). Rev. ital. Paleont., Bologna, 2 : 352-356, pi. 6, 9 figs. 
Viereck, H. L. 1912. Contributions to our Knowledge of Bees and Ichneumon-flies, in- 
cluding twenty-one new genera and fifty-seven new species of Ichneumon-flies. Proc. U.S. 

natn. Mus., Washington, 42 : 613-648, 2 figs. 
Werner, F. G. 1969. In Anderson, R. Y. & Kirkland, D. W., Terrestrial insects of the 

Rita Blanca lake deposits. Mem. geol. Soc. Am., Washington, 113 : 123-130, pi. 19. 
Weyenbergh, H. 1869. Sur les insectes fossiles du calcaire lithographique de la Baviere, 

que se trouvent au Musee Teyler. Archs Mus. Teyler, Haarlem, 2 : 247-294, pis 34-37. 
Wheeler, W. M. 1926. Les societes d'insectes ; leur origine, leur evolution. 468 pp., 61 figs. 

Paris. [English translation 1928.] 



fWHiTiNG, P. W. 1935. Sex determination in bees and wasps. /. Hered., Washington, 

26 : 229. 
WiLLE, A. 1959. A new fossil stingless bee (Meliponini) from the amber of Chiapas, Mexico. 

/. Paleont., Tulsa, 33 : 849-852, pi. 119, figs 1-4. 
1964. Notes on a primitive stingless bee, Trigona (Nogueirapis) mirandula. Revta Biol. 

trop., San Jose, 12 : 117- 151, 21 figs. 
& Chandler, L. C. 1964. A new stingless bee from the Tertiary amber of the Dominican 

Republic (Hymenoptera ; Meliponini). Revta Biol, trop., San Jose, 12 : 187-195, 5 figs. 
Zeuner, F. E. 1931. Die Insektenfauna des Bottinger Marmors. Fortschr. Geol. Palaeont., 

Berlin, 9 : 247-406, pis 1-19. 

1938. Die Insektenfauna des Mainzer Hydrobien-kalks. Palaeont. Z., Berlin, 20 : 104- 

151, pis 13-17- 

1944- Fossil Honey-Bees. Brit. Bee-Keep. Ass., London : 10. 

1951- A Discussion of Time-rates in Evolution. Proc. Linn. Soc. Lond. 162 : 124-130. 


New taxonomic names and the page numbers of the principal references are printed in bold 

Aix-en-Provence 250 

amber, see Baltic, Burmese, Sicilian 

Andrena 165-6, 167-9, 171, 175 

clavula 166, 167 

grandipes 166, 167 

hypolitha 166, 167 

lagopus 167 

percontusa 167 

primaeva 167-8, 183 ; pi. i, fig. 2 

sepulta 166, 167 

tertiaria 202 

wrisleyi 166 

spp. indet. 168-9 
Andrenidae 165, 166-71 
Andreninae 165, 166-70 
Anthidiini 178, 256 
Anthidium 178, 179 

curvatum 179 

exhumatum 179 

mortuum 178-9, 193 

scudderi 179 
Anthophora 193, 194, 198, 204-5 

effossa 200, 202 

melfordi 194 ; pi. i, fig. i. 
Anthophoridae 187, 188-99 

gen. et spp. indet. 200 

gen. et sp. indet. 256 
Anthophorinae 192, 193-9, 256 
Anthophorini 192, 193, 194-9 
Anthophorites 193, 194-5, 196-9 

gaudryi 195, 197 

longaeva [longaevus) 195-6 

mellona 193-5, 196-7 

melona 196 

parietaria 195 
thoracica {thoracicus) 195 
titania 197-8 
tonsa {tonsus) 198 
veterana {veteranus) 198-9 
ants 202, 219 
Apiaria dubia 186 

veterana 255 
Apidae 206, 207-54 
gen. et spp. indet. 254 
incertae sedis 250-4 
Apini 206, 225, 226-49 
Apis 177, 202, 213, 226-8, 230, 232, 
240-2, 243, 244-51 ; see Synapis 
acervorum 194 
adamitica 176-7, 186 
amalthea 216-7 
aquitaniensis 250-1 
armhrusteri 244-6, 247-8 
scharmanni 246, 247 
scheeri 246, 247 
scheuthlei 246, 247-8, 249 
seemanni 248 
zeuneri 248 
bicornis 183 
catanensis 257 

avolii 257 
centuncularis 182 
cucurbitina 188 
cuenoti 240 

dormitans 184, 241, 251 
dorsata 237, 243, 247, 249 
floreaz^T, 245, 249 
helvola 166 




iridiae 243 

malvae 193 

manicata 178 

meliponoides 225, 227, 229, 231 

melisuga 201, 248-9 

mellifera 225, 237, 241, 243-8, 249 ; pi. 4, 
fig- 6 
indica 243 

mellifica 239, 243, 249 

oligocenica [oligocaenica) 240-2 

palmnickenensis 233 

quadricincta 163 

yw/a 183 

styriaca 251 

terrestris 209 

truncorum 180 

violacea 189 

sp. 247 
Apoidea 161 et seqq. 

fossils erroneously classified as 202-3, 257 

gen. et spp. incertae sedis 201-2, 256-7 
Armbruster, L. 158 
auricle 158 
Axestotrigona erythra 217 

togoensis 217 

Baltic amber 159-60, 166, 168, 172-5, 200, 

207-8, 214, 228-39, 252-7 
Bauckhorn, H. 160 
Bedford, R. 205 
bees' nests 203, 256 
Biareolina 165, 167, 170 

neglect a 165, 167 
Blattidium beroldingianum 255 
Bombini 157, 206, 209 

Bombus igo, 201, 209, 210-3, 226-7, 232, 
237. 254 

ahavus 210-1, 212-3 '< P^- 2, fig. 2 

antiquus 252-3 

carbonarius 252, 254 

conservatus 255 

crassipes 209-10 

florissantensis 201, 212, 213 

grandaevus 201, 212, 253 

junnei 189-90 

lapidarius 213 

muscorum 252 

proavus 212-3 

pusillus 252, 254 

terrestris 209 
Bombusoides mengei 253 
Bottingen 160, 245 
Burmese amber 160, 223 

Burrow, Col. H. 161 
Bursey, Mr 229 

Calyptapis 201, 209, 212 

florissantensis 209, 212 
Camoins 160 
Catania 257 
Celliforma 203, 204-6 

bedfordi 204-5 ; pi. i, figs 7, 8 

favosites 203, 206 

germanica 204 

«W(ia 204 

septata 205 ; pi. i, fig. 9 ; pi. 2, fig. i 

spirifer 203-4 

sp. indet. 205 
Centris clavipes 221 

cornuta 176 
Ceratina 166, 188 

cucurbitina 188 

disrupta 188 
Ceratinini 188 
Cereste 240 

Chalcobombus 206-7, 208, 214, 227-8, 232 
group 206 

hirsutus 207 

humilis 206-7, 208 

martialis 208 
Chalicodoma 182-3 
Chiavone 201 
Clavicera i88 
Cockerell, T. D. A. 159 
cockroaches 255 
CoUetidae 161-2, 255 
copal 160, 217-21, 249 
Corent 195-6 
Crematogaster 219 
Ctenoplectra 172 
Ctenoplectrella 172, 173 

dentata 172 

splendens 255 

viridiceps 172, 173 
Ctenoplectrinae 172, 173-4 
Cyr tapis 163 

anomalus 163 

Dasypoda 171, 175 
depstum 158, 230 
Diandrena 170 
Dianthidium 179, 180 

curvatum 179 

tertiarum 180 
Dominica 257 



East African copal i6o 

Electrapis 206, 225-7, 228-39, 254 ; see 
Protobombus, Roussyana 

apoides 226, 227-9, 237 

meliponoides 225, 227, 229-31, 233, 237 ; 
pi. 3, figs 5, 6 

minuta 257 

tornquisti 231, 237 
Eucera 178-9, 201, 212, 253 

antennata 193 

mortua 178-9, 193 
Eucerini 192-3 
Euglossa 237 
Euglossini 206 

Florissant 161, 163 et seqq. 
Formicinae 202 
Fraas, O. 158 

Gabbro 201, 249 

Gard 193 

Glyptapis 172, 173, 174 

fuscula 173 

mirabilis 173-4 

neglecta 174 

reducta 174 

reticula 174 

reticulata 174 

Halictidae 162, 163-5 
Halictinae 162-3, 164-6 
Halictoides maurus 170 
Halictus 163, 167-8, 171, 175, 198 

florissantellus 164 

miocenicus 164 

ruissatelensis 164 

schemppi 164-5 

scudderiellus 164 
Haplomelitta 174 
Hauff, B. 159 
Hauffapis 239, 243-4 

scharmanni 244, 246 

scheeri 244, 247 
gallauni 247 
rahdei 247 

scheuthlei 243-4, 247 
seemanni 247-8 
zeunevi 247-8 
Heer, O. 161 
Heriades 180, 181 -2 

bowditchi 181 

halictinus 181 

laminarum 181, 182 

mersatus 181 

mildredae 181-2 

priscus 182 

5a;\^05M5 182 

truncorum 180 
Hermann, P. 159 
Hesperapis rhodoceratus 170 
Hukong Valley 160, 223 
Hylaeus albilabris 188 
Hypotrigona 218, 219-21 

bottegoi 218 

dominicana 256 

eocenica 256 

gribodoi 218-21 ; pi. 2, figs 5-7 

Klebs, R. 159 
Krantz, F. 160 
Krottensee 209 

larval chambers 203-6, 256 
Lerida 155 

Lestrimelitta 206, 213, 217 
Libellulapis 170, 171 

antiquorum 170 

wilmattae 170, 171 
Liotrigona bottegoi 218 
Lithandrena 165, 169 

antiquorum 169 

saxorum 169 
Lithanthidium 180 

pertriste 180 
Lithoblatta lithophila 255 
Lithophaga nuda 204 
Lithurge 176, 177-8 

adamitica 176-8 

cornutum 176 
Lithurginae 176, 177-8 
Lithurgus 176-7 

adamiticus 177 
Luke Thomas Collection 160 

Macrocera 193 

Mainz 159 

Manning, Dr F. J., obituary 156 

Megachile 179-80, 182, 183 

amaguensis 182 

centuncularis 182 

praedicta 183 

sp. indet. 183 



Megachilidae 172, 175, 176-87 

gen. et sp. indet. 186 
Megachilinae 176, 178, 179-86 
Megachilini 180, 181 -6 
Megilla pilipes 194 
Melectini 192, 199 
Melipona 213, 224, 254 

togoensis 217 
Meliponini 206, 213, 214-25 
Meliponorytes 216, 223 

devictus 222-4 

sicula 216, 224 

succini 216-7, 221 
Meliponula 217 

boccandei 217 
Melissodes 212 
Melitta 175 

tricincta 175 

miliar di 175 
Melittidae 171, 172-5 
Melittinae 174-5 
Menat 256 

Mesoblattina Uthophila 255 
Mexico 257 

Michener, C. D. 156, 161, 192 
Musca Uthophila 255 

native bee 257 
Naucoris carinata 255 
Nogueirapis silacea 257 
Nomada 167 
Nomia 169 

Oeningen 159, 161, 177, 185-6, 190, 196-8, 

201, 210 
Orthoptera 255 
Osmia 167, 180, 183, 184-6, 202, 205 

antiqua 183, 184-5 

bicornis 183 

carbonum 184, 253 

dormitans 251 

sp. ? dubia 183, 185-7 

immortua 183, 186 

kirbyana 183, 202-3 

nigra 183, 185-6 ; pi. i, figs 5, 6 

primaeva 167, 183 

spp. indet. 186, 256 

Panurginae 170, 171 

Panurgus 170 

Parandrena andrenoides 170 

Parschlug 251 
Pelandrena 165, 169, 170 

re duct a 169, 170 
Perdita 170 
Piski 202 

Podalirius 194, 204 
Polistes 203 

kirbyanus 202-3 
Probombus hirsutus 256 
Prohalictus 163 

schemppi 163-4 
Protobombus 227, 231-2, 233 

indecisus 231, 232-3 

tristellus 233 
Protomelecta 199 

brevipennis 199 

Radoboj 159, 195, 253 

Randecker Maar 158, 164, 244, 246-8 

Rohwer, S. A. 159 

Ross, H. 157 

Rott am Siebengebirge 157, 178, 184 

200, 202, 240, 242-3, 251 
Roussy, L. 160 
Roussyana 227, 233, 234-9 

palmnickenensis 226, 233-6, 237, 
pi. 3, figs I, 2 

proava 228, 236-9 ; pi. 3, figs 3, 4 


Scheele Collection 160 

Scheer, E. 159 

Schempp, K. 159 

Scheuthle, W. 158 

Siberia 182 

Sicilian amber 158, 224 

Simjovel 257 

Snodgrass, R. E. 156 

Solnhofen 255 

Sophrobombus 206, 213-4, 215-6 

fatalis 213, 214-6 ; pi. 2, figs 3, 4 
South Australia 160 
Sphecodes 164 
Sphecoidea 155-6 
Spokane 212 
Statz, G. 160 
Swinhoe, R. J. C. 160 
Synapis 232, 240, 241-3, 249 

cuenoti 240, 241 

henshawi 237, 240-3 

dormiens 241-2, 251 ; pi. 4, figs 1-5 
kaschkei 240, 243 



Tetragona 216, 221, 222-4 

elongata 221 

iridipennis 222-4 ; pi. 2, figs 8, 9 

succini 221-2, 224 
Tetragonula 222 
Tetralonia 193 

antennata 193 

berlandi 192, 193 

duvaucelii 193 
tibial comb, rake, spurs 157 
Timon-David Collection 160 
Tosi, A. 158 

Trigona 214, 216-7, 218-26, 236-7, 254 ; 
see Hypotrigona, Nogueirapis, Tetragona 

amalthea 2x6 -j 

braunsi 218 

bottegoi 218, 220 

devictus 222 

dominicana 256 

elongata 221 

eocenica 256 

gi'>'^Am 217-8 
togoensis 217 

gribodoi 218 

iridipennis 222 

laeviceps 222-4 

sicula 224-5 
togoensis 217-8 
Trinidad 255 

Uruguay 205-6 
auroranormai 205-6 

wasp 183, 202 

nest 203 
Wheeler, Dr W. M. 159 
Wickham, Professor H. F. 161 
wing venation 157 

Xilocopa 189 

Xylocopa 189, 190-2, 210-1 

abavus 210 

friesei 189 

hydrobiae 189 

jurinei 189-90 

senilis 189, 190-2 ; pi. i, figs 3, 4 

violacea 189 
Xylocopinae 187-8, 189-92 
Xylocopini 188-9, 190-2 

Zeuner, Professor F. E., obituary 156 

Professor F. E. Zeuner, 

Ph.D., D.Sc, F.S.A., F.G.S., F.Z.S.f 
F. J. Manning, Ph.D.f 

S. F. Morris 

Department of Palaeontology 

British Museum (Natural History) 

Cromwell Road 

London SW7 5BD 

Accepted for publication 2 December 1975 

Anthophora melfordi Cockerell (p. 194) 
Fig. I. Holotype, BM(NH) 1. 9259. Oligocene, Florissant. X2-5. 

Andrena primaeva Cockerell (p. 167) 
Fig. 2. Holotype, ETH Zurich SN. 202. Miocene, Oeningen. X5. 

Xylocopa senilis Heer (p. 190) 
Figs 3, 4. ETH Ziirich 21. Miocene, Oeningen. x 2, x 2-2. 

Osmia nigra sp. nov. (p. 185) 
Fig. 5. Holotype, ETH Ziirich PI. I. 719. Head and thorax. Miocene, Oeningen. x 4-5. 
Fig. 6. Holotype, ETH Ziirich PI. L 719. Abdomen. Miocene, Oeningen. x 4-5. 

Celliforma bedfordi sp. nov. (p. 204) 
Fig. 7. Holotype, BM(NH) In. 3 1434. Lateral view. Pleistocene or Sub-Recent, Venus 
Bay, South Australia, x 0-9. 

Fig. 8. Holotype, interior view. Horizon and locality as Fig. 7. x 2-0. 

Celliforma septata sp. nov. (p. 205 ; see also PI. 2, fig. i) 
Fig. 9. Holotype, BM(NH) In. 3 1433. Horizon and locahty as Fig. 7. x i. 

Bull. Br. Mils. nat. Hist. (Geol.) 27, 3 


>' -'-^ 

;-, '>,1»S*4''''*'3^***- 

Celliforma septata sp. nov. (p. 205 ; see also PI. i, fig. 9) 
Fig. I. Paratype, BM(NH) In. 31435. Longitudinal section. Pleistocene or Sub-Recent, 
Venus Bay, South Australia. x i. 

Bombus abavus Heer (p. 210) 
Fig. 2. ETH Zurich PI. L 808. Miocene, Oeningen. x 3-3. 

Sophrobombus fatalis Cockerell (p. 214) 

Fig. 3. Hamburg, Scheele Collection 1225. Eocene, Baltic amber. x 7-5. 
Fig. 4. Hamburg, Scheele Collection 1225. Eocene, Baltic amber. x 8. 

Trigona (Hypotrigona) gribodoi Magretti (p. 218) 
Fig. 5. BM(NH) In. 38984. Pleistocene copal, East Africa. x 10. 
Fig. 6. BM(NH) In. 38984. Pleistocene copal. East Africa. x 12. 
Fig. 7. BM(NH) 5851 1. Pleistocene copal. East Africa. x 10. 

Trigona (Tetragona) iridipennis Smith (p. 222) 
Fig. 8. BM(NH) In. 43809. ? Pleistocene, Burmese amber, Hukong Valley, Burma. x 11. 
Fig. 9. BM(NH) In. 20702. ? Pleistocene, Burmese amber, Hukong Valley, Burma. x 13. 

Bull. Br. Mils. nat. Hist. (Geol.) 27, 3 










r ^B 

Sfc^ -^^'Z^^hj 




r 9 

Electrapis (Roussyana) palmickenensis (Roussy) (p. 233) 
Figs i, 2. Hamburg, Scheele Collection 218. Eocene, Baltic amber. Fig. i, x 11 ; Fig. 2, 

Electrapis (Roussyana) proava (Menge) (p. 236) 
Fig. 3. Lectotype, BM(NH) In. 43592. Eocene, Baltic amber. x 5. 
Fig. 4. Paralectotype, BM(NH) In. 18757. Eocene, Baltic amber. x 4-5. 

Electrapis (Electrapis) meliponoides (Buttel-Reepen) (p. 229) 
Figs 5, 6. Hamburg, Scheele Collection 238. Eocene, Baltic amber. x 7-5. 

Bull. Br. Mus. nat. Hist. (Geol.) 27, 3 


Apis (Synapis) henshawi dormiens subsp. nov. (p. 241) 
Fig. I. Statz Collection. Oligocene, Rott. x 4. 
Fig. 2. BM(NH) In.36655. Oligocene, Rott. x 5. 
Fig. 3. Statz Collection. Oligocene, Rott. x 4. 
Fig. 4. BM(NH) 59634. Oligocene, Rott. x 2-6. 
Fig. 5. Statz Collection. Oligocene, Rott. x 4. 

Apis (Apis) mellifera Linnaeus (p. 249) 
Fig. 6. BM(NH) 58576. Pleistocene copal, East Africa. x 4. 

Bull. By. Mils. nat. Hist. (Geol.) 27, 3 

u«- ^' » ,■ 



' -1 ■ 

\. . 








1. Cox, L. R. Jurassic Bivalvia and Gastropoda from Tanganjdka and Kenya. 
Pp. 213 ; 30 Plates ; 2 Text-figures. 1965. out of print. 

2. El-Naggar, Z. R. Stratigraphy and Planktonic Foraminifera of the Upper 
Cretaceous — Lower Tertiary Succession in the Esna-Idfu Region, Nile Valley, 
Egj^t, U.A.R. Pp. 291 ; 23 Plates ; 18 Text-figures. 1966. £11. 

3. Davey, R. J., DowNiE, C, Sarjeant, W. a. S. & Williams, G. L. Studies on 
Mesozoic and Cainozoic Dinoflagellate Cysts. Pp. 248 ; 28 Plates ; 64 Text- 
figures. 1966. £8.20. 

3. Appendix. Davey, R. J., Downie, C, Sarjeant, W. A. S. & Williams, G. L. 
Appendix to Studies on Mesozoic and Cainozoic Dinoflagellate Cysts. Pp. 24. 
1969. 95p. 

4. Elliott, G. F. Permian to Palaeocene Calcareous Algae (Dasycladaceae) of 
the Middle East. Pp. iii ; 24 Plates ; 16 Text-figures. 1968. out of print. 

5. Rhodes, F. H. T., Austin, R. L. & Druce, E. C. British Avonian (Carboni- 
ferous) Conodont faunas, and their value in local and continental correlation. 
Pp- 313 \ 31 Plates ; 92 Text-figures. 1969. £13.10. 

6. Childs, a. Upper Jurassic Rhynchonellid Brachiopods from Northwestern 
Europe. Pp. 119 ; 12 Plates ; 40 Text-figures. 1969. £5.25. 

7. Goody, P. C. The relationships of certain Upper Cretaceous Teleosts with 
special reference to the Myctophoids. Pp. 255 ; 102 Text-figures. 1969. 


8. Owen, H. G. Middle Albian Stratigraphy in the Anglo-Paris Basin. Pp. 164 ; 
3 Plates ; 52 Text-figures. 1971. £7.20. 

9. SiDDiQUi, Q. A. Early Tertiary Ostracoda of the family Trachyleberididae 
from West Pakistan. Pp. 98 ; 42 Plates ; 7 Text-figures. 1971. £9.60. 

10. Forey, p. L. a revision of the elopiform fishes, fossil and Recent. Pp. 222 ; 
92 Text-figures. 1973. ;fii.35- 

11. Williams, A. Ordovician Brachiopoda from the Shelve District, Shropshire. 
Pp. 163 ; 28 Plates ; 11 Text-figures ; no Tables. 1974. £12.80. 

PrinUd in Great Britain by John Wright and Sons Ltd. at The Stonebridge Press, Bristol BS4 SNU 

' GtWERAL ^\ 

2 7 OCT 1976 

^, UBRARY ^^./ 


GEOLOGY Vol. 27 No. 4 

LONDON: 1976 



2 7 OCT 1976 


Pp 271-345 ; 12 Plates ; 12 Text-figures 

GEOLOGY Vol. 27, No. 4 

LONDON: 1976 


(natural history), instituted in 1949, is 
issued in five series corresponding to the Scientific 
Departments of the Museum, and an Historical series. 

Parts will appear at irregular intervals as they 
become ready. Volumes will contain about three or 
four hundred pages, and will not necessarily be 
completed within one calendar year. 

In 1965 a separate supplementary series of longer 
papers was instituted, numbered serially for each 

These papers form Vol. 27, No. 4^ of the Geological 
[Palaeontological) series. The abbreviated titles of 
periodicals cited follow those of the World List of 
Scientific Periodicals. 

World List abbreviation : 
Bull. Br. Mus. nat. Hist. (Geol. 

ISSN 0007-1471 

© Trustees of the British Museum (Natural History), 1976 


Issued 28 October, 1976 Price £7-95 


Fusella M'Coy 1844, a problematic brachiopod genus from the Lower 

Carboniferous. By C. H. C. Brunton & A. Rissone .... 275 

The Jurassic Chaetetid, Blastochaetetes hathonicus J. C. Fischer, in England. 
By G. F. Elliott 285 

Coprolites of Ptilophyllum cuticles from the Middle Jurassic of north 
Yorkshire. By C. R. Hill 289 

A new fossil freshwater crab from the Ngorora Formation (Miocene) of 

Kenya. By S. F. Morris . 295 

Cyclothyris (Cretaceous Brachiopoda) from California. By E. F. Owen . 301 

Cranial material of Oligocene and Miocene flamingos : with a description of 

a new species from Africa. By C. J. O. Harrison & C. A. Walker . 305 

A new fossil pelican from Olduvai. By C. J. O. Harrison & C. A. Walker 315 

Chelidonocephalus trilobite fauna from the Cambrian of Iran, By R. A. 


Index to Vol. 27 no. 4 341 






The genus Fusella is redescribed and assigned to the subfamily Strophopleurinae. Related 
or similar species are discussed. 


The genus name Fusella, first published in 1844 by M'Coy for small transversely 
fusiform spiriferide brachiopods, has been poorly known and ill-used for about one 
hundred and thirty years. We redescribe the type specimen of the type species, 
F. fusiformis (Phillips), and other conspecific and congeneric material in the hope 
of establishing the genus Fusella on a more stable basis. Silicified specimens of 
F. rhomhoidea (Phillips) allow the description of interiors believed to be closely 
comparable to those of F. fusiformis. The genus is formally redescribed and we 
discuss both its position within the Spiriferacea and those species which, in the past, 
have been assigned to Fusella. 


Superfamily SPIRIFERACEA King 1846 


Subfamily STROPHOPLEURINAE Carter 1974 

Diagnosis (emended). Small to medium very transverse Mucrospiriferidae with 
simple lateral costae and lirate, subimbricate micro-ornamentation. Fold and 
sulcus commonly non-costate, with or without median rib and groove ; sulcus 
bounding ribs commonly accentuated, as are corresponding dorsal grooves. Ventral 
interarea large and denticulate. Dental plates short or buried by shell thickening. 
Dorsal sockets small and closely set ; cardinal process commonly medially supported 
by short ridge ; shell substance impunctuate. 

Discussion. Recently Carter (1974) proposed a classification of the Spiriferidae 
which involved the erection of the Strophopleurinae. Into this subfamily he placed 
' Stfophopleura Stainbrook, 1947 ; Alispirifer Campbell, 1961 ; Acuminothyris 
Roberts, 1963 ; Voiseyella Roberts, 1964 { = Amesopleura Carter, 1967) ; ? Eleuthero- 
komma Crickmay, 1950 ; ? Pterospirifer Dunbar, 1955 ; ? Celsifornix Carter 
(1974 : 677). This subfamily corresponds partially with a new taxon of the authors' 
which was in script form at the time of Carter's 1974 publication. We accept 

Bull. Br. Mus. nat. Hist. (Geol.) 27, 4 : 275-284 Issued 28 October 1976 


Carter's subfamily but emend its taxonomic position and, importantly for the 
purpose of this paper, add the genus Fusella as a firm member of the subfamily. 
We believe that Alispirifer Campbell doubtfully belongs here but would suggest 
the inclusion of Brachythyrina Frederiks 1929 and Paeckelmanella Likharev 1934. 
Roberts (1971) placed Voiseyella in the Mucrospiriferidae but the above grouping 
within the Strophopleurinae removes Fusella and Brachythyrina from the 
Spiriferidae. Alispirifer, Paeckelmanella and Pterospirifer are removed from the 
Licharewiinae and Eleutherokomma from the Acrospiriferinae of the Treatise 
(Williams et al. 1965) classification. 

Ivanova (1972 : 315) proposed the family Paeckelmanellidae for Paeckelmanella, 
Alispirifer, Spiriferinaella and Pterospirifer. The family was placed, with some 
reservation, in the Syringothyridacea, within the suborder Spiriferidina, but 
neither a full diagnosis nor a discussion was provided. Carter's subfamily, here 
used, partially equates with Ivanova's family, as can be seen from the generic 
constituents. Ivanova, Carter and we ourselves all agree that the taxon containing 
Paeckelmanella - and we believe Fusella - belongs within the Spiriferidina, but at 
family levels of classification there is no agreement ; Ivanova placed the mucro- 
spiriferids within the new suborder Delthyrididina whereas most previous authors 
placed them in the Spiriferidina. 

Genus FUSELLA M'Coy 1844 

Type species. Spirifera fusiformis Phillips 1836 : 210 ; pi. 9, figs 10, 11. By 
original designation of M'Coy (1844 : 132). 

Diagnosis. Small (commonly less than 30 mm wide) strongly transverse shells 
with pointed extremities. High, variably concave, denticulate ventral interarea 
extending full width of shell. Lateral profile subcircular. Ventral sulcus bordered 
by pair of prominent ribs, dorsal fold variably developed. Lateral ribbing weak 
to moderately developed. Dental plates close together within sulcus, subparallel 
and with callosity filling apex of delthyrium in large specimens. Crural bases 
converge to valve floor posteriorly. SheU impunctate. 

Discussion. Although M'Coy (1844) was reasonably precise, for that date, in his 
description of Fusella, the name has been ill-used ever since. This is because there 
is only one well-known specimen of the type species F. fusiformis in existence and 
from this it is impossible to learn any detailed information about the internal 
morphology. It is this species which M'Coy specified as the type of his new 
'subgenus' Fusella, characterized as foUows. 

'Shell elongate transversely, fusiform, cardinal area wide, much curved ; beaks 
incurved. This group would embrace these little Spirifers of the mountain 
limestone which have a perfectly fusiform outline, the depth being equal to the 
length, and the sides cylindrical ; the cardinal area is extremely wide in proportion 
to their size and is always hollowed or much curved, thus contrasting with the 
narrow, flat area of the typical Spirifers while the strongly incurved beaks 
distinguish them from the Cyrtiae. It would include the S. hicarinata, S. 
rhomboides, &c. &c.' (1844 : 132). 


M'Coy's description stressed the incurved nature of the ventral umbo, which leads 
to the behef that he had not seen Phillips' actual specimen of F. fusiformis in which 
the ventral interarea is almost flat, only being concave medially, close to the umbo. 
The umbo projects beyond the hinge line by no more than 1-5 mm ; Waterhouse 
(1970 : 3) is in error in writing that the 'umbo extended 2-5 mm beyond the hinge', 
probably because he took his measurement from his pubhshed illustrations which 
are at almost twice the stated magnification. M'Coy's description may therefore 
have been influenced by specimens from the Cork area of Ireland best assigned to 
F. rhomboidea (PhiUips). This suggestion is further supported by the size quoted 
by M'Coy for F. fusiformis, viz. 'length four lines, width one inch three lines, depth 
four lines^' (1844 • 132), a width which is somewhat greater than that of the type 
specimen of F. fusiformis. 

A review of the confused use of the name Fusella was published by Waterhouse 
(1970). He redescribed the type specimen, from the Gilbertson Collection in the 
British Museum (Natural History), and compared it with various other species in 
an attempt to suggest its affinities. He concluded that Unispirifer Campbell 1957, 
with which Fusella has sometimes been synon3nnized, was distinctive and we agree 
with this view. Waterhouse thought that the shell substance of F. fusiformis was 
punctate, leading him to discard species such as Spirifer rhomboidea PhiUips as being 
closely related, but to the conclusion that Fusella was 'probably related to members 
of the Sjnringothyrididae' (1970 : 6). Both optical and scanning electron-microscope 
studies of the type specimen and second undoubted specimen of F. fusiformis show 
that the shell is not endopunctate but quite normal for impunctate spiriferaceans 
(PL I, figs 20, 21). 

Thomas (1971) is one of the latest of several palaeontologists to say that it seemed 
inadvisable to use the generic name Fusella until the type specimen was adequately 
known from topotypic material. Had the name fallen from use this would be a 
sensible suggestion but in view of its continued appearance in the literature, 
commonly quite incorrectly, it is desirable to further Waterhouse's contention that 
it should become a well-known genus in its own right. Following the Russian lead 
when Ivanova (i960) placed 5. tornacensis de Koninck within Fusella some 
palaeontologists, such as Carter (1967), have placed species in Fusella which differ 
widely from F, fusiformis. Within their concept the genus is relatively less wide, 
very much longer and has a strongly uniplicate anterior commissure. Carter (1971) 
described the genus Mirifusella, said to be 'most similar to Fusella M'Coy', but in 
fact differing considerably in outhne and internal features. We have, therefore, a 
situation in which some palaeontologists advise the suppression of the name Fusella 
and others use this name, at times quite incorrectly. Because of this confusion it 
is desirable to correct the use of Fusella to the best of our ability, even if this is done 
without resort to additional genuine topotypic material. In the collections of the 
British Museum (Natural History) there exists one specimen clearly conspecific with 
the type specimen of F. fusiformis. This second specimen is in the Davidson 
Collection (B 7379) and came from Dovedale, Derbyshire (PI. i, figs 5-7). The 
ventral umbo has been broken from this specimen and it is possible to see that the 

1 A line or ligne is one twelfth of an inch ( = 2-1 167 mm). 


dental plates did not extend anterodorsally to support completely the teeth and 
delthyrial margin near the hinge line. Unfortunately searches in the Derbyshire 
or Bolland and Clitheroe areas have failed to reveal additional specimens. The 
information on internal morphology of Fusella is, therefore, mainly based upon the 
silicified Fermanagh specimens collected by Brunton and assigned to F. rhomhoidea 
(Phillips), a species believed to be closely related to F.fusiformis and which originally 
also came from Bolland (PL i, figs 13-19). 

In the last twelve years two genera have been proposed which are probably synony- 
mous with each other and also are closely related to Fusella. These are Voiseyella 
Roberts 1964, with type species Strophopleura anterosa Campbell 1957 from the 
Lower Carboniferous of the Werria Basin, New South Wales, Australia, and 
Amesopleura Carter 1967, with type species Spirifer novamexicana Miller 1881 from 
the Lower Carboniferous (Osagian) of New Mexico. Both these authors thought 
that Spirifer mundula Rowley 1893 should probably be placed in their genus. 
Subsequently Roberts (1971) has placed ^meso^/^«*ra into synonymy with Voiseyella. 
Having inspected Roberts' 1964 material from the Greenhills area of New South 
Wales and Carter's 1967 material from the Lake Valley region of New Mexico we 
agree with this sjmonymy. The question then arises as to the relationship of 
Voiseyella with Fusella. Neither Roberts nor Carter compared their genera with 
Fusella, although the former (1971) discussed the relationship of Fusella with 
Unispirifer and the latter placed his new species llanoensis (1967) within his concept 
of Fusella. These genera are clearly quite closely related ; their dimensions, outlines 
and profiles are very similar, as is the form of ribbing. The most important differ- 
ence, and that which prevents the synonymy of Voiseyella within Fusella, is that 
the dental plates of Voiseyella diverge from the umbo following the ribs bordering 
the sulcus whilst those of Fusella are unusual in remaining subparallel within the 
confines of the ventral sulcus (PI. i, fig. 19) ; they do not follow the borders of the 
sulcus, as in many spiriferaceans. (The dental plates of V. novamexicana also follow 
the ribs bordering the ventral sulcus.) 

In view of the varied use of the name Fusella, and despite the redescription by 
Waterhouse (1970), it seems desirable to provide a description of F. fusiformis 
based upon the type specimen and second specimen in the Davidson Collection. 

Fusella fusiformis (Phillips) 

PI. I, figs 5-12 

1836 Spirifera fusiformis Phillips : 217 ; pi. 9, figs 10, 11. 

1849 Spirifera fusiformis Phillips ; Brown : 108 ; pi. 51, figs 4, 5. 

1858 Spirifera fusiformis Phillips ; Davidson : 56 ; pi. 13, figs 15, 15a. 

1970 Fusella fusiformis (Phillips) ; Waterhouse : 3 ; figs lA-F. 

Type specimen. The single specimen described and figured by Phillips (1836) 
in the Gilbertson Collection, British Museum (Natural History), B 249, from 
Bolland, Yorkshire. 

Dimensions. Width (incomplete) 22-6 mm. Mid-point to ear of more complete 
side 13-8 mm. Length 8-2 mm. Thickness 8-2 mm. Angle of sulcus 33°. An 
estimate of the complete width of the shell is about 28 mm. 


Diagnosis. Small transversely narrowly rhombic shells as long as thick and 
about three and a third times as wide as long. High ventral interarea with only 
slightly projecting umbo. Ribbing weakly developed and dorsal median fold 
prominent only anteriorly ; dorsal umbo medially sulcate. Dental plates short and 
subparallel, diverging anteriorly less than the angle of sulcus. 

Description. The tip of the right side of the shell is missing and sediment 
obscures two thirds of the interarea. The external surface has been deeply exfoliated 
in patches, especially on the ventral valve. Elsewhere the shell is somewhat eroded 
and on only one small area in the ventral sulcus is primary shell preserved. Thus 
details of external ornamentation cannot be given, nor is it possible to count the 
number of ribs with certainty other than on the left half of the dorsal valve where 
there are ten or eleven simple costae. Within the more deeply exfoliated secondary 
fibrous shell of the ventral valve short radially arranged traces of mantle canals are 
visible. Growth lines are clearly seen, especially on the dorsal valve (about 4 per mm 
at the side of the fold) and indicate a fusiform shape throughout ontogeny ; they 
became more prominent anteriorly. The ventral interarea is vertically grooved by 
flexures in the secondary fibres (PI. i, figs 10, 21) which formed a delicate denticula- 
tion of the hinge line at the inner surface. (When covered by primary shell this 
feature would not be visible.) The ventral sulcus is prominent but the expected 
dorsal fold is virtually lacking although its position is marked by a pair of bordering 
ribs more prominent than the others. There is a dorsal median depression which 
becomes shallower and less well defined anteriorly from the umbo. The anterior 
commissure is uniplicate. No internal structures can be seen in the specimen 
although the exfoliation of the ventral umbo shows the secondary fibres to be bent 
along lines interpreted as the bases of the dental plates. If this is correct, it can be 
said that the dental plates are positioned on the borders of the ventral sulcus at a 
distance of 2 mm from the umbo but anteriorly remain subparallel and thus within 
the diverging sulcus. (This is the same as is seen in the Fermanagh silicified material 
assigned to Fusella.) 

The second specimen, from Dovedale (B 7379, PL i, figs 5-7), is smaller than the 
holotype and in that it too has lost its right tip and ventral umbo it is less well 
preserved (half width 11-3 mm, length c. 6-3 mm, thickness 6-i mm). The dorsal 
fold is slightly developed anteriorly, resulting in a rather more prominent uniplication 
of the anterior commissure than in the holotype. The broken ventral umbo allows 
it to be seen that the dental plates are short for they did not reach anteriorly to the 
plane of fracture. It is impossible to measure the angle of divergence of the ventral 
sulcus but an estimate of the delthyrial angle, based on the remaining dorsal half 
of the interarea, is 30°. As in the holotype, the dorsal umbo is unusually shaped 
with a shallow V-shaped groove about i mm long terminating anteriorly on the 
pair of large ribs bordering the fold. Inspection of the visible surfaces shows that 
the shell structure of both specimens is fibrous with no sign of endopunctation. 

Discussion. It is unfortunate that F. fusiformis is a rare species in rocks of low 
to mid Visean age. If it were not for the second conspecific specimen from Dovedale 
we might have considered the type specimen to be a freak. Indeed it is possible 



that the presence of a reasonably developed dorsal fold is more characteristic ; 
this feature is better developed on the Dovedale specimen and it is unusual for a 
uniplicate spiriferacean not to have a better developed fold. Growth lines show that 
even at a dorsal valve length of 2-5 mm the commissure was uniplicate. The 
amplitude of the uniplication is 3-0 mm on the type specimen and 3-2 mm on the 
Dovedale specimen. The growth lines also show that these shells grew with a 
fusiform shape virtually throughout their postembryonic ontogeny ; a growth line 
about I mm from the dorsal umbo indicates that the valve width at that stage 
was 4 mm. 

Fig. I. Camera-lucida drawings of silicified dorsal valve interiors of F. rhomboidea (Phillips) 
from the Visean of Co. Fermanagh, Ireland. A, B, dorsal and posterodorsal views of a 
nearly fully developed valve ; C, dorsal view of part of a fully developed valve ; D, dorsal 
view of a juvenile valve at twice the magnification. Note the posteriorly narrow sockets 
which, in adult shells, remained functional only anteriorly. Cross shading denotes 
broken shell material, a.s. - adductor muscle scar ; b.c. - broken stump of the crus ; 
c. - crus ; c.p. - cardinal process ; d.i. - dorsal interarea ; d.u. - dorsal umbo ; m.r. - 
median ridge (dividing the adductor scars) ; s. - socket ; s.r. - socket ridge, which 
merges dorsally with the crural base. (See also PI. i, fig. 18.) 





Fig. 2. Camera-lucida drawings of a silicified ventral valve of F. rhomhoidea (Phillips) 
from the Visean of Co. Fermanagh, Ireland. A, posterior view ; B, lateral view ; 
C, ventrolateral view. a.d. - apex of the delthyrium, filled by secondary shell material ; 
d.p. - dental plate ; d.r. - dental ridges bordering the interior surfaces of the edges of 
the delthyrium. Anteroventrally these ridges are supported by the dental plates which 
buttress across the ventral shell cavity, i.v.i. - internal surface of the ventral interarea ; 
s.t. - secondary shell thickening between the dental plates ; t. — tooth ; v.s. - ventral 
sulcus. (See also PL i, fig. 19.) 

In the British Isles the species most closely resembling F. fusiformis is F. 
rhomhoidea (Phillips), the type specimen of which also came from BoUand (PI. i, 
figs 13-17). This species is a constituent of the silicified brachiopod faunas being 
studied by Brunton (1966, 1968) from Ireland but as yet has not been redescribed 
except briefly by Brunton & Champion (1974). Other than F. rhomhoidea there are 
several species mentioned by Waterhouse (1970), some of which he believed may 
belong to Fusella. It was his mistaken belief that F. fusiformis was endopunctate 
which led him to discard species like rhomhoidea, convoluta (Phillips) and trigonalis 
(Martin) as being closely related. For the same reason Waterhouse related Fusella 
to Syringothyris , in particular some specimens believed to be S. cuspidatus (J. 
Sowerby). He figured (1970 : fig. 2, A-F) a specimen named 'Spirifer cuspidatus 
(Phillips)' from Treak Cliff, Derbyshire, in the British Museum (Natural History) 
collections (BB 40831). This specimen belongs neither to the species cuspidatus 
(first described by Martin in 1809 but ascribed to J. Sowerby (1816) by Muir-Wood 


when she selected the lectotype in 1951) nor, since it is impunctate, to Syringothyris 
which includes only endopunctate species. The outline, even for a young specimen 
of cuspidatus, is too transverse, and furthermore true cuspidatus does not have a 
sulcate fold or denticulate ventral interarea such as that of the Treak Cliff specimen. 
Of the Gilbertson Collection specimens (B 297) from BoUand, labelled as 5. cuspidatus 
and discussed by Waterhouse, only the largest three are true endopunctate Syringo- 
thyris cuspidatus. The fourth is poorly preserved and of doubtful affinity but the 
fifth is impunctate and like the Treak Cliff specimen BB 40831. These specimens 
should probably be assigned to the Strophopleurinae but do not accord with any 
presently described British species. Their dental plates are more like those of 
Voiseyella than Fusella. 

Spirifer distans Sowerby is another species discussed by Waterhouse (1970) in the 
belief that it is closely related to F. fusiformis. Although some specimens assigned 
to distans may resemble F. fusiformis and despite true 5. distans being impunctate, 
we do not believe it to be a Fusella species. The Sowerby type specimen of S. distans, 
from near Dublin, Ireland (B 61009), differs from F. fusiformis in external shape 
and a second Irish specimen (B 7664), more complete than the type refigured by 
Davidson (1858 : pi. 8, figs 5-8), is illustrated here for comparison (PI. i, figs 1-4). 
The dental plates of this species diverge, following the borders of the ventral sulcus, 
and there is an apically complete arched delthyrial plate. 

Another species belonging in the subfamily but remaining difficult to assign to a 
genus, through a lack of internal information, is Spirifer roemerianus de Koninck, 
from the Tournaisian of Belgium. The same is true for the species Spirifer 
triangularis J. de C. Sowerby, placed in Fusella by Muir-Wood in 195 1. This species 
is larger than F. rhomboidea and differs in having a high carinate fold and prominent 
ventral median rib in the sulcus. Spirifer convoluta Phillips is another extremely 
transverse species but it reaches much larger dimensions (at least 80 mm wide) than 
F. fusiformis. Its interior is unknown so the generic assignment is doubtful, but 
if it were to be included within Fusella the diagnosis of that genus would require 
emendation to include species at this size. Spirifer hicarinata M'Coy was one of the 
species M'Coy mentioned originally as being in Fusella. Other than by M'Coy's 
description and incomplete figure, 5. hicarinata is virtually unknown ; the type 
specimen seems to be lost and it is unwise to continue using the name. 

In conclusion, therefore, we assign Fusella to the Strophopleurinae and in addition 
to the type species, F. fusiformis (Phillips), we include F. rhomboidea (Phillips) and 
doubtfully F. trigonalis (Martin), F. triangularis (J. de C. Sowerby), F. roemerianus 
(de Koninck) and F. convoluta (Phillips) within this genus. Strophopleura probably 
evolved from the Mucrospiriferidae in the Upper Devonian and gave rise to the 
Tournaisian Voiseyella and Acuminothyris and to the Visean Fusella ; from this the 
Carboniferous to Lower Permian genus Brachythyrina may have evolved, and also 
the northern Permian genera Paeckelmanella and Pterospirifer . 


Brown, T. 1849. Illustrations of the fossil conchology of Great Britain and Ireland, with 
descriptions and localities of all the species, viii +273 pp., 117 pis. London. 



Brunton, C. H. C. 1966. Silicified productoids from the Visean of County Fermanagh. 
Bull. Br. Mus. nat. Hist. (Geol.), London, 12 ; 173-243, pis 1-19. 

1968. Silicified brachiopods from the Visean of County Fermanagh (II). Bull. Br. Mus. 

nat. Hist. (Geol.), London, 16 : 1-70, pis 1-9. 

& Champion, C. 1974. A Lower Carboniferous brachiopod fauna from the Manifold 

Valley, Staffordshire. Palaeontology, London, 17 (4) : 811-840, pis 107-111. 

Carter, J. L. 1967. Mississippian brachiopods from the Chappel Limestone of central Texas. 
Bull. Am. Paleont., Ithaca, 53 : 253-488, pis 13-45. 

1971. New early Mississippian silicified brachiopods from central Iowa. Smithson. 

Contr. Paleobiol., Washington, 3 : 245-255, pis 1-2. 

1974- New genera of spiriferid and brachythyridid brachiopods. /. Paleont., Tulsa, 

48 (4) : 674-696, pis 1-4. 

Davidson, T. 1858. A monograph of British Carboniferous brachiopods, 2 (5) i . Palaeontogr. 

Soc. {Monogr.), London : 1-48, pis 1-8. 
IvANOVA, E. A. i960. Spiriferida. In Orlov, Y. A. (ed.). Principles of Palaeontology, (7) 

Bryozoa and Brachiopoda : 264-280. Moscow (Akad. Nauk SSSR), [In Russian.] 

1972. Main features of Spiriferid evolution (Brachiopoda). Paleont. J., Washington, 

6 (3) : 309-320. [Translated from the Russian.] 

M'CoY, F. 1844. A synopsis of the characters of the Carboniferous Limestone fossils of Ireland. 
207 pp., 29 pis. Dublin. 

MuiR-WooD, H. M. 1951. The Brachiopoda of Martin's "Petrifiicata Derbiensia". Ann. 
Mag. nat. Hist., London, (12) 4 : 97-118, pis 3-6. 

Phillips, J. 1836. Illustrations of the geology of Yorkshire. Part II. The Mountain Lime- 
stone district. 253 pp., 25 pis. London. 

Roberts, J. 1964. Lower Carboniferous brachiopods from Greenhills, New South Wales. 
/. geol. Soc. Aust., Adelaide, 11 (2) : 173-194, pis 1-6. 

1 97 1. Devonian and Carboniferous brachiopods from the Bonaparte Gulf Basin, North- 
western Australia. Bull. Bur. Miner. Resour. Geol. Geophys. Aust., Melbourne, 122 : 1-399, 
pis 1-59. 

Thomas, G. A. 1971. Carboniferous and early Permian brachiopods from Western and 

Northern Australia. Bull. Bur. Miner. Resour. Geol. Geophys. Aust., Melbourne, 56 : 1-272, 

pis 1-31. 
Waterhouse, J. B. 1970. The Lower Carboniferous brachiopod genus Fusella M'Coy 1844. 

Life Sci. Occ. Pap. R. Ont. Mus., Toronto, 15 : 1-12, pis i, 2. 
Williams, A. et al. 1965. Brachiopoda. In Moore, R. C. (ed.). Treatise on Invertebrate 

Paleontology, H. 927 pp., 746 figs. Kansas. 

C. H. C. Brunton, Ph.D. 


Department of Palaeontology 
British Museum (Natural History) 
Cromwell Road 
London SW7 5BD 

Accepted for publication 9 February 1976 


The specimens are all housed in the BM(NH) and those in Figs 1-19 were coated with 
ammonium chloride sublimate before being photographed. The prints are not retouched. 

Spirifer distans J. de C. Sowerby 

Figs 1-4. Visean of Millicent, Co. Kildare, Ireland. Viewed anteriorly, posteriorly, dorsally 
and from the left side, x i. B 7664. 

Fusella fusifortnis (PhilHps) 
Figs 5-7. Visean of Dovedale, Derbyshire. Viewed anteriorly, posteriorly and dorsally. 
The arrow on Fig. 7 indicates the region illustrated in Fig. 20. x 2. B 7379. Davidson 

Figs 8-12. Holotype, figured by Phillips, from Bolland, Yorkshire. Viewed anteriorly, 
dorsally, posteriorly (the arrow indicates the region enlarged in Fig. 21), ventrally and from 
the left side, x 1-5. B 249. Gilbertson Collection. 

Fig. 20. Exfoliated standard secondary fibres from the position marked on Fig. 7. The 
posterior dorsal margin is to the top and the mid-line to the right. Scanning electron 
micrograph , x 2 1 o . 

Fig. 21. Exfoliated secondary fibres of the ventral interarea of the holotype (see Fig. 10) 
showing the flexures resulting in a fine denticulation at the commissure. Scanning electron 
micrograph, x 140. 

Fusella rhomboidea (Phillips) 

Figs 13-17. Lectotype, figured by Phillips, from Bolland, Yorkshire. Viewed ventrally, 
anteriorly, posteriorly, dorsally and from the right side, x i ; Figs 15, 17x2. B 236. 
Gilbertson Collection. 

Figs 18-19. Silicified specimens from the Upper Visean of the Sillees river, Co. Fermanagh, 
Ireland (see Brunton 1966 for locality details). Fig. 18, mature dorsal valve interior (see 
also Fig. lA-D, p. 280), x 4. BB 61611. Fig. 19, young ventral valve interior looking 
posteriorly (see Fig. 2A-C, p. 281), x 3. BB 61612. 

Bull. Br. Mus. nat. Hist. (Geol.) 27, 4 





Three English occurrences are hsted of Blastochaetetes bathonicus Fischer, a species abundant 
in the French Bathonian east of the Paris Basin. It is suggested that their location, rarity and 
growth-forms may be due to migration, south of the Ardennes -London landmass, into less 
favourable conditions in the Cots wold area of southern England. 

The post-Palaeozoic chaetetids were monographed by J. C. Fischer (1970). He 
discussed the structures and classification of these somewhat problematic calcified 
radial-tubular colonial organisms in detail, and compared them mth bryozoa, 
tabulates and stromatoporoids, concluding that they were referable to the Hydrozoa 
(Coelenterata ; Cnidaria), His species Blastochaetetes bathonicus (Fischer 1965, 
1970) occurs commonly in the Middle Bathonian of the Aisne and Ardennes in 
northern France, and is also recorded from Haute-Marne and the Cote-d'Or ; all 
these areas are in the Jurassic outcrop east of the Paris Basin. In Aisne and 
Ardennes the fossil is abundant in the 'Calcaires pseudo-oohthiques superieurs', in 
which it forms characteristic banks or beds at some levels. Commonly it is 
accompanied by algal oncolites in a coarse-grained calcareous rock, suggesting that 
it flourished during hfe in agitated waters (Fischer 1969a, 1969b). 

In England it appears to be a rare fossil, and only three occurrences are known to 
me, all Bathonian ; two are in Gloucestershire, and one near Bath (Elliott 1972, 
1974). These records are now given in detail. 

1. Great Oolite, probably near top of White Limestone. Tiltup's End, Nailsworth, 
Glos. For description of this old quarry, now overgrown, see Witchell (1886) 
and Woodward (1894 : 270). The specimen is in the British Museum (Natural 
History) Dept. of Palaeontology, registered number R 49314, Witchell Collection. 

2. Great OoUte, probably near top of White Limestone. Limekiln Pit, Royal 
Agricultural CoUege, Cirencester, Glos. Another old exposure (Woodward 
1894:282). BM(NH) Dept. Palaeont. reg. no. R 49313, Hinde Collection 
(? ex Tomes Collection). Plate 2, figs 1-3. 

3. Great Oolite, Upper Rags, Bed 3 of Elliott (1974). Bathampton Down, east of 
Bath, Avon (formerly Somerset). BM(NH) Dept. Palaeont. reg. no. R 49603, 
G. F. EUiott CoUection. Plate 2, fig. 4. 

In thin section all three specimens show the characteristic internal structures 
and approximate dimensions as given in full detail by Fischer (1970 : 179) ; the 
amber-coloured tube-walls with fibrous 'jet d'eau' or feather-structure are beautifuUy 
preserved. The external growth forms, however, are very different. The Tiltup's 

Bull. Br. Mus. nat. Hist. (Geol.) 27, 4 : 285-288 Issued 28 October 1976 


End specimen is a compact hemispherical growth exactly like that given as typical 
for the species by Fischer (1970 : fig. 20a, b), but is a small example (diameter 30 mm, 
as opposed to an average of 40 mm, maximum 120 mm, in the French material), 
No matrix remains. The Cirencester example is larger (approximate diameter 
55 mm), higher-domed, and in section is seen to be grown round a coral nucleus 
of 15 mm diameter. It is interesting that this nucleus is almost completely altered 
to clear crystalline calcite, traces of identifying coral-structure surviving only at the 
margins, in contrast to the excellent preservation of the enveloping Blastochaetetes. 
The whole specimen is in a white pseudoolitic rock. Finally the Bathampton 
example occurs as thin irregular spreads, possibly crusts, of less than 10 mm thick- 
ness ; they are in an ochreous organic-rubble limestone with brachiopods, sponges, 
molluscs, echinoderms and bryozoa, whole and broken. The rock is interpreted as a 
current-accumulated deposit (Elliott 1974). This 'spreading' Blastochaetetes is 
regarded as a growth form only, for the Tiltup's End and Cirencester examples both 
contain old external surfaces heavily overgrown by later developments of the same 

The obvious and shortest migration-route in Bathonian times, from the colonies 
of the Aisne- Ardennes where the species flourished, was westwards in suitable 
facies south of the Ardennes-London landmass, and so to the clear-water calcareous 
site of the present Cotswolds in England. The areas to both north and south of 
this were in different facies. The species did not become abundant in England, and 
presumably did not survive into the changed conditions indicated by the overlying 
Forest Marble facies (Upper Bathonian). The recent 'Tentative reconstruction 
of the palaeogeography of part of southern England at the end of White Limestone 
times' (Palmer & Jenkins 1975 : fig. 6) indicates the Cotswold area as offshore- 
current marine. The numerous small facies-changes seen there in the actual rocks 
at about this level, as well as the rare varied growth forms of B. hathonicus, may well 
be evidence of less favourable conditions for the species than those which existed 
in France. 

Note. Hartman & Goreau (1972), after a comparison with living Sclerospongiae, 
regarded the Chaetetida as extinct sclerosponges. 


Elliott, G. F. 1972. Cretacicrusta gen. nov., a possible alga from the English Cretaceous. 
Palaeontology, London, 15 : 501-503, pis 100, loi. 

1974- Note on the palaeoecology of a Great Oolite fossil-bed at Bath (English Jurassic). 

Proc. Geol. Ass., London, 85 : 43-48. 

Fischer, J. C. 1965. Decouverte d'un niveau-repere a "Chaetetidae" dans le Bathonien 
moyen ardennais. C. r. hebd. SSanc. Acad. Sci., Paris 260 : 6641-6643. 

1969a. Geologie, paleontologie et paleoecologie du Bathonien au sud-ouest du Massif 

ardennais. Mem. Mus. natn. Hist. nat. Paris (n.s. C) 20. 320 pp., 20 pis. 

1969b. Sur I'existence d'un cycle biostratigraphique dans le Bathonien au sud-ouest du 

Massif ardennais (Note preliminaire) . Bull. Soc. geol. Fr., Paris, (7) 10 : 303-307. 

1970. Revision et essai de classification des Chaetetidae (Cnidaria) post-Paleozoiques. 

Annls Paldont. (Invert.), Paris, 56 : 149-233, 6 pis. 


Hartman, W. D. & GoREAU, T. F. 1972. Ceratoporella (Porifera : Sclerospongiae) and the 

Chaetetid "Corals". Trans. Conn. Acad. Arts Sci., New Haven, 44 : 131 -148, 26 figs. 
Palmer, T. J. & Jenkyns, H. C. 1975. A carbonate island barrier from the Great Oolite 

(Middle Jurassic) of central England. Sedimentology , Oxford &c., 22 : 125-135. 
Witchell, E. 1886. On the Forest Marble and Upper Beds of the Great Oohte, between 

Nailsworth and Wotton-under-Edge. Proc. Cotteswold Nat. Fid Club, Gloucester, 8 : 267- 

280, pis 4, 5. 
Woodward, H. B. 1894. The Jurassic Rocks of Britain, IV. The Lower Oolitic rocks of 

England (Yorkshire excepted). Mem. geol. Surv. U.K., London, 628 pp. 

G. F. Elliott, D.Sc, Ph.D., F.G.S.. F.L.S. 

Department of Palaeontology 

British Museum (Natural History) 

Cromwell Road 

London SW7 5BD Accepted for publication 9 February 1976 

Blastochaetetes bathonicus J. C. Fischer 
Fig. I. Solid specimen, 'cauliflower-head' growth embedded in matrix, x 2. M. Jurassic, 
Great Oolite (Bathonian) ; Limekiln Pit, Royal Agricultural College, Cirencester, Glos. 
BM(NH) Dept. Palaeont., reg. no. R 49313. 

Fig. 2. The same specimen ; thin section, x 15, showing radial growth with old outer surface 
overgrown by a differently orientated second growth. 

Fig. 3. The same specimen ; portion of outer surface, x 15. 

Fig. 4. Thin section of a spreading or laminar growth, x 15. M. Jurassic, Great Oolite 
(Bathonian) ; Bathampton Down, near Bath, Avon. BM{NH) Dept. Palaeont., reg. no. 
R 49603. 

Bull. By. Mus. nat. Hist. (Geol.) 27, 4 







Large coprolites from the Roseberry Topping plant bed (Middle Jurassic) consist mainly 
of Ptilophyllum fragments. Their significance is discussed. 


The specimens discussed here were collected over 50 years ago by Dr H. Hamshaw 
Thomas, though so far as I know he never described them. They were found in 
siltstones of Lower Deltaic age which form the upper layers of the Roseberry Topping 
plant bed (Thomas 1913). Besides being much the largest of the half dozen or so 
coprolites now known from the Yorkshire Middle Jurassic they have yielded an 
entirely different flora from those previously investigated. 

The material is in the Collection of the Palaeontology Dept. of the British 
Museum (Natural History), and consists of six blocks labelled V 58510 and an 
additional one labelled V 58510a. Those labelled V 58510 look as if they once 
formed part of a single larger block which broke up in collecting, and judged on the 
six fragments this original block must have been large, having more than 250 pellets 
covering an area of about i m^. As the pellets occur on several bedding planes, 
occupying about 5 mm thickness of the cross-bedded matrix, I assume that their 
burial was rapid. Otherwise I would expect them to have become widely dispersed, 
whereas they are in fact somewhat clustered into random groups, very like fresh 
droppings of a modern herbivore such as a rabbit. Indeed, much like freshly 
dropped rabbit pellets the round or oval outlines of the fossil ones sometimes show a 
constriction to a point on the side from which they were extruded, and fine striations, 
presumably caused by the anal sphincter, may be seen radiating from this point 
onto the impression surfaces. The clarity of these markings in the fossil doubtless 
indicates that the fresh pellets were of a firm consistency and also of a uniform, 
possibly gelatinous, texture. • 

The diameter of the pellets ranges from 8 to 18 mm, though they are mostly 
10-13 iiin^- Where present the substance, which is composed of compressed cuticle 
fragments and finely divided organic detritus, is a dull black colour. It is about 
0-5 mm thick, thinning to the edges, and the lack of compression border suggests 
that the pellets were perhaps somewhat flattened before their final burial (com- 
pression theory of Walton 1936). The fact that adjacent pellets sometimes overlap 
one another may equally indicate that they were originally rather flattened (Harris 

Bull. Br. Mus. nat. Hist. (Geol.) 27, 4 : 289-294 Issued 28 October 1976 

2 go 


Fig. 3. Coprolite, chiefly of Ptilophyllum pectinoides (Phillips) Phillips, from Roseberry 
Topping, north Yorkshire. V 58510, x i. Drawn on a photograph. 


1946). However they could well overlap if, as in the pelleted form of sheep dung, 
they were sometimes produced adhering together in masses. 

V 58510a may have come from a different horizon of the Roseberry Topping 
siltstone as the colour of the matrix is rather darker. The pellets have a similar 
form to those of V 58510 but they are uniformly smaller, about half the size, and 
many more of them overlap their neighbours. As, however, they yield an almost 
identical flora to those of V 58510 I imagine that they may well have been produced 
by the same kind of animal, but possibly an individual having a smaller anus ; by 
analogy a young rabbit produces much smaller pellets than a fully grown one. 

Of the abundant cuticles yielded by Schulze- Ammonia maceration over 99% 
of the determinable ones are fragments from leaves and petioles of Ptilophyllum 
pectinoides (Phillips) Phillips. Only a few scraps from other species were seen, 
chiefly a specifically indeterminable Solenites or Czekanowskia and a Bennettitalean 
cuticle resembling Nilssoniopteris vittata (Brongn.) Florin (though possibly also 
referable to certain species of Anomozamites or Pterophyllum) . There are in addition 
several well-preserved micropylar and interseminal scale cuticles identical with those 
of Williamsonia hildae Harris and W. leckenhyi Nathorst. 

The size of the cuticle fragments reaches about i mm x 3 mm, though it is usually 
I mm^ or smaUer. The substance of the pellets, however, shows marked fracturing 
into rectangular blocks and this, which presumably occurred after deposition, may 
well have reduced the size of some of the cuticle fragments. 


Coprolites from the Yorkshire Deltaic beds are rare and informative fossils. 
Other than occasional insects, fish skeletons and reptilian footprints they provide the 
only evidence so far known for the activity of vertebrate animals in the delta. The 
examples previously described, collected from the Gristhorpe Bed and at Beast 
Cliff, have been discussed by Harris (1946, 1951, 1956, 1964) and Harris et al. (1974). 
None of them, however, has pellets even as much as half the size of those in the present 
specimens and they have yielded entirely different floras, chiefly composed of 
comparatively delicate leaf and seed cuticles or pollen : mainly Caytonia, Solenites, 
Androstrobus, and thin Bennettitalean cuticles. In contrast to these the leaf of the 
Roseberry specimens, Ptilophyllum pectinoides, had a robust cuticle. Frequently 
also this leaf was resinous, as is indicated both in hand specimens and in the coprolite 
by the internal casts of mesophyll cells sometimes seen adhering in patches to the 
upper cuticle (PI. 3, figs i, 2, 3 ; Harris 1949). I conclude that the Roseberry 
herbivore was a rather larger animal than those which provided Harris's specimens, 
perhaps the size of a sheep or large rabbit. Indeed the present pellets are remarkably 
similar to the pelleted form of sheep's dung. The animal clearly could utilize the 
rather harsh resinous leaves of Ptilophyllum, though it did not digest the resin to 
any noticeable extent. 

The nature of the herbivore is, of course, unknown, though presumably it was a 
reptile, such as an ornithopod dinosaur, or possibly a mammal. If a mammal then 
the pellets seem larger than would be expected from any mammals of Middle 


Jurassic age so far described in the literature. Gut dimensions, however, though 
generally related to size, do not display a direct mathematical proportionality to it, 
and the analogy to the size of a sheep is approximate. 

The intimate association, within the coprolite, of the flower Williamsonia with the 
leaf Ptilophyllum is striking. These organs are also seen associated together as 
hand specimens in the normal flora at Roseberry and on this basis, with additional 
evidence from morphologically intermediate organs, Harris (1969) has attributed 
them to the same plant. In this respect it seems that the coprolite provides a 
remarkably faithful representation of the normal Roseberry flora. Assuming that 
the animal fed on living plants the evidence thus reinforces Harris's restoration. 

The overall floristic composition of the dung is also like that of the normal flora 
at certain layers in the upper part of the siltstone, and this is interesting in view of 
the complex factors, such as fragmentation and sorting, which normally operate 
during the deposition, as at Roseberry, of terrestrial plants in stream channel 
deposits. As it is natural to assume that the herbivore fed on living vegetation the 
facts suggest that the fragmentary debris which constitutes the normal fossil flora 
might sometimes be a remarkably straightforward representation of plant 
communities then growing in the vicinity. They may also indicate that the plants 
grew fairly close to Roseberry, for if the animal had strayed far one might expect 
the dung to yield at least a few species not normally found there. 

There is, however, a simpler explanation. The animal may possibly have been a 
scavenger, browsing on the Ptilophyllum debris which was being transported in the 
deltaic streams and ultimately deposited in great abundance at Roseberry and other 
localities. If this is right the dung would be expected to show fragments of species 
we already know from the normal Roseberry flora, but it would not necessarily 
indicate anything about plant communities in the living vegetation other than that 
their debris could be palatable to an unknown animal. 

Unfortunately there is insufficient evidence to make a certain choice between these 
explanations, though so far as I know there are no modern vertebrates which would 
normally prefer a diet of rotting leaves to fresh ones. For this reason the first 
explanation seems perhaps nearer the truth. 


Harris, T. M. 1946. Notes on the Jurassic Flora of Yorkshire, 19-21. Ann. Mag. nat. Hist., 
London, (11) 12 : 357-378. 

1949. Notes on the Jurassic Flora of Yorkshire, 43-45. Ann. Mag. nat. Hist., London, 


1951- Notes on the Jurassic Flora of Yorkshire, 49-51. Ann. Mag. nat. Hist., London, 


1956. The investigation of a fossil plant. Proc. R. Instn Gt Br., London, 36 (163) : i-ii. 

1964. The Yorkshire Jurassic Flora, 2 : Caytoniales, Cycadales and Pteridosperms. viii + 

191 pp., 7 pis. Brit. Mus. (Nat. Hist.), London. 

1969. The Yorkshire Jurassic Flora, 3: Bennettitales. vi + i86 pp., 7 pis. Brit. Mus. 

(Nat. Hist.), London. 

MiLLiNGTON, W. & Miller, J. 1974. The Yorkshire Jurassic Flora, 4 : Ginkgoales and 

Czekanowskiales. viii+150 pp., 8 pis. Brit. Mus. (Nat. Hist.), London. 



Thomas, H. H. 1913. The Jurassic plant beds of Roseberry Topping. Naturalist, London, 
1913 : 198-200. 

Walton, J. 1936. On the factors which influence the external form of fossil plants ; with 
descriptions of the foliage of some species of the Palaeozoic Equisetalean genus Annularia 
Sternberg. Phil. Trans. R. Soc, London, (B) 226 (535) : 219-237. 

C. R. Hill, B.Sc, Ph.D. 

Department of Palaeontology 
British Museum (Natural History) 
Cromwell Road 
London SW7 5BD 

Accepted for publication 9 February 1976 

Ptilophyllum pectinoides (Phillips) Phillips 

Figs i, 2, 3. Upper cuticle prepared by Schulze maceration of a hand specimen found at 
Hasty Bank. Fig. i, V 58591, light micrograph, x 160. Fig. 2, inner surface of cuticle; 
scanning electron micrograph showing resinous casts inside the 'cutinized' walls of palisade 
mesophyll cells, x 336. Fig. 3, same, x 1040. 

Fig. 4. Coprolite, chiefly of Ptilophyllum pectinoides, from Roseberry Topping. V 58510, x i. 
Photo : J. V. Brown. 

Bull. Br. Mus. nat. Hist. (Geol.) 27, 






A new species of freshwater crab, Potamonautes {Lirrangopotamonautes) tugenensis, is de- 
scribed from the Miocene of the Tugen Hills, Kenya. It is compared with Recent species of 
P. {Orthopotamonautes) and P. [Lirrangopotamonautes). 


Dr Martin Pickford of Bedford College, London, whilst collecting fossil mammals 
from the Lake Baringo area of northern Kenya, collected a number of claystone 
nodules from the northern end of the Tugen Hills, about 20 miles (32 km) north-west 
of Lake Baringo. They came from the two neighbouring localities of Kamwina, 
Kabarsero area (Pickford locality 2/72) and Kalimale area (Pickford locality 2/68). 
The Kamwina locality is a lake shoreline deposit with diatomaceous shales and the 
Kalimale section is a relict river channel. Kamwina is Member C and Kalimale 
Member D of the Ngorora Formation of Miocene age, approximately 11-12x10^ 
years (Bishop & Chapman 1970, Bishop, Chapman, Hill & Miller 1971). The 
Kalimale locality has 5delded only isolated limb segments. For the latest assessment 
of the chronostratigraphy of the region, Berggren & van Couvering (1974) should 
be consulted. 

The nodules collected by Dr Pickford contain freshwater crabs and ostracods. 
The brittle nodules are not laminated in any way so that attempts to break them 
open by hammer resulted in the shattering of the nodule. An attempt to develop 
the nodules by acid treatment was unsuccessful. It remained to split each one with 
a rock splitter, which was less damaging than the hammer but nevertheless reduced 
it to a number of fragments that had to be glued back together. If there was any 
preferred plane of splitting it was through the abdomen and along the smoother 
subhepatic and pterygostomial areas. Even though the specimens are in nodules 
they are crushed. Fortunately Dr Pickford had collected one specimen, the holotype, 
which had weathered free of matrix and was not badly crushed, and also some 
isolated segments of limbs. Altogether about 50 nodules were collected that 
contained crab remains. The limb fragments from Kalimale are not sufficiently 
different to suggest that a second species is present, but limb fragments are not very 
diagnostic. A modern freshwater crab species would probably not occupy both 
such diverse habitats as river channels and lake shores, since this would involve 
different water speeds, temperature and bottom conditions. Hence it is possible 

Bull. Br. Mus. nat. Hist. (Geol.) 27, 4 : 295-300 Issued 28 October 1976 


that a second species is present, but that problem can only be resolved by more and 
better material from Kalimale. 


Section BRACHYRHYNCHA Borradaile 1907 
Superfamily XANTHOIDEA Dana 1851 

Family POTAMIDAE Ortmann 1896 
Genus POTAMONAUTES Macleay 1838 

Type species. Thelphusa perlata H. Milne-Edwards 1837, by subsequent 
designation of Rathbun 1905 : 159. 


Type species. Potamon [Potamonautes) lirrangensis Rathbun 1904, by original 
designation of Bott 1955 : 264. 

Potamonautes (Lirrangopotamonautes) tugenensis sp. nov. 
PL 4, figs 1-6 

Diagnosis. A Lirrangopotamonautes with strongly produced front and medially 
divided urogastric area, single spines on the carpus and on the otherwise smooth 

Material, One more or less complete carapace, 50 distorted and fragmentary 
carapaces and some isolated chelipeds. The holotype and most of the paratypes 
are in the National Museum of Kenya, Nairobi. The remaining paratypes (nos 
In 61517-24) are in the British Museum (Natural History), London. The material 
comes from two Miocene localities in the Ngorora Formation, at Kamwina and 
Kalimale about 20 miles (32 km) NW of Lake Baringo, Tugen Hills, Kenya. 

Description. Carapace one quarter wider than long at its widest point about 
one third back. The front is moderately long and strongly produced with only a 
slight ventral deflection ; it is smooth and occupies about one third of the anterior 
carapace width. The anterior margin is smooth and strongly sinuous with a 
thickened edge. On the holotype one ocular peduncle is preserved. The post- 
frontal crest is strong, overhangs the front and is nearly straight with a slight 
deflection towards the front at the mid-line ; it meets the anterolateral margin at 
an obtuse angle. The postfrontal crest and anterolateral margins are smooth or 
slightly beaded. The margins of the orbits are entire, thickened and smooth. 
The single extraorbital spine is well developed. The median furrows join at or only 
just posterior to the postfrontal crest and pass forward on to the frontal area, 
cutting the frontal area into two. The lateral margins are convex anteriorly and 
may be slightly concave posteriorly. 

The anterolateral marginal ridge passes up on to the dorsal carapace just behind 
the widest part of the carapace. The posterolateral marginal ridge starts below the 


point at which the anterolateral marginal passes on to the dorsal carapace, and goes 
in an anteroventral direction. 

The hepatic areas are inflated and traversed by two rows of pits. The line of the 
pits curves proximally across the branchial area to join the cervical groove opposite 
the widest part of the mesogastric area. The urogastric area is clearly delimited by 
the cervical groove and the furrow between the uro- and metagastric areas. The 
urogastric area is divided by a longitudinal furrow. The proximal walls of the 
cervical groove are deeply hachured on the internal mould (and probably also on 
the shell itself). The cervical groove is deep medially but becomes shallower 
anterolaterally. The rhomboidal branchial areas are inflated and covered with 
feeble rugosities. 

The ventral surface is poorly preserved - usually only the first four thoracic 
sternites or hemisternites are preserved. No abdomina have been found except for 
one poorly preserved female, of which the segments appear simple and not fused. 
The third maxilliped is reasonably well preserved although no merus has yet been 
found. The ischium is broad and subrectangular ; it bears a groove parallel to the 
inner margin and about one third in. The exopodite is about a third of the width 
of the ischium and tapers anteriorly. The posterior inner angles of the ischia close 
against a knob-like anterior extension of the first thoracic sternite, which has the 
shape of a 'parson's nose'^. The lateral flanges of the anterior extension have two 
small swellings, sometimes extended into ridges, one on each flange. Although 
corresponding depressions have not been found on the underside of the ischia, they 
appear to have a press-stud effect presumably to locate accurately the third 
maxilliped when it is closed. The fingers of the larger, right, cheliped are approxi- 
mately the same length as the palm and are gently curved towards the mid-line. 
The fingers are grooved laterally and bear large teeth. There is virtually no gape 
between the fingers, which are opposed throughout their length. The smaller 
cheliped is similar to the larger one except that the palm is narrower and the teeth 
of the fingers fine with an occasional larger tooth. The smooth carpus bears a 
single spine at the distal end of the inner marginal ridge. The ridge is sharp and 
strongly elevated. The merus also bears a single spine at the distal end of its inner 
marginal ridge but the outer marginal ridge is strong and smooth or may possibly 
have a few coarse granules. 


Very few species of freshwater crabs have been found fossil and this is the first that 
can be assigned to a Recent genus. The form of the postfrontal crest and the shape 
and position of the epibranchial angle clearly indicate that it belongs to the Recent 
African genus Potamonautes. Potamonautes was split by Bott (1955) into fifteen 
subgenera, to a large extent on the form of the gonopods. However there are 
sufficient carapace differences to allow the fossils to be compared with the two 
subgenera Lirrangopotamonautes and Orthopotamonautes . Bott (1955 : 310) erected 
four new genera for fossil species of potamid crabs, but stated that on the characters 

^ Pygostyle of domestic fowl. 


of the carapace, which alone is usually preserved, the fossil species could be compared 
with Recent species. The closest species to the present fossils are the living P. 
{Orthopotamonautes) depressus (Krauss) and P. {Lirrangopotamonautes) lirrangensis 
(Rathbun). Both these are restricted geographically to central and east Africa at 
the present time. 0. depressus has a weakly produced or even deflected front 
whereas L. lirrangensis has a strongly produced front similar to L. tugenensis. 
However L. lirrangensis has a denticulated anterolateral border and three rows of 
denticles on the merus of the cheliped. The widest part of the carapace of 0. 
depressus lies just in front of the anterior third, whilst that of L. tugenensis lies just 

The many species of Potamonautes are distributed through central and southern 
Africa, especially in the tropical belts and the Rift Valley system. The majority 
have discontinuous distributions at the present day in the cooler water of the 
montane regions where they are found in many habitats from rapid-flowing rivers 
to shallow trickles or even standing water (Williams 1968) . But some species groups, 
e.g. the 0. johnstoni group (Williams 1968) to which L. tugenensis probably belongs, 
prefer the slow-moving warmer waters of the lakes and especially the coastal belts. 
The Kamwina fossils appear to come from lacustrine deposits in which the water 
could have been warm and slow-moving. The discontinuous occurrences of the 
Recent species appear to reflect a residuum of previous land forms. The single 
fossil species does not allow any reconstruction of previous drainage patterns, but a 
number of occurrences might give some clue both to the evolution of the drainage 
pattern and to the interrelationship of the many species of freshwater crabs from 
east Africa. Dr Keith Bannister is at present trying to unravel the drainage 
pattern of east Africa by studying the interrelationships of various species of the 
large fish Barbus which are also distributed through many separated bodies of water 
(personal communication) . 

Although L. tugenensis is the first fossil freshwater crab from Africa that can be 
assigned with any certainty to a modern genus, Joleaud & Hsu (1935 : 108 ; fig. 11) 
published a specimen, described as Totamonide gen. nov.', from Cenomanian 
terrestrial deposits of Tanout, Niger (Furon 1935 : 35), which is difficult to assess 
from the outline figure. Unfortunately the figured specimen cannot be found at 
present in Paris. In the form of the postfrontal crest it could certainly belong to 
the Potamidae near Potamonautes but the two lateral crests do not conform to any 
Recent genus. 


Berggren, W. a. & VAN CouvERiNG, J. A. 1974- The Late Neogene : Biostratigraphy, 
geochronology and paleoclimatology of the last 15 million years in marine and continental 
sequences. Palaeogeogr. Palaeoclimat. Palaeoecol., Amsterdam, 16 : 1-216. 

Bishop, W. W. & Chapman, G. R. 1970. Early Pliocene sediments and fossils from the 
northern Kenya Rift Valley. Nature, Lond. 226 : 914-918. 

, Hill, A. & Miller, J. A. 1971. Succession of Cainozoic vertebrate assemblages 

from the northern Kenya Rift Valley. Nature, Lond. 228 : 389-394. 

BoRRADAiLE, L. A. 1907. On the classification of the Decapoda. Ann. Mag. nat. Hist., 
London, (7) 19 : 457-486. 



BoTT, R. 1955. Die Susswasserkrabben von Afrika und ihre Stammesgeschichte. Annls 

Mus. r. Congo Beige, Terverun, C. Zoologie (III, III) 1 (3) : 209-352, 30 pis, 103 figs. 
Dana, J. D. 1851. On the Classification of the Cancroidea. Am. J. Set., New Haven, (2) 

12 : 121-131. 
FuRON, R. 1935. Le Cretace et le Tertiaire du Sahara soudonais. Archs Mus. natn. Hist. 

flat. Paris (6) 13 : 1-96, pis 1-7. 
JOLEAUD, L. & Hsu T.-Y. 1935. Crustaces decapodes du Cretace de Tanout (Damergou 

Niger fran9ais). Archs Mus. natn. Hist. nat. Paris (6) 13 : 99-110, 11 figs. 
Macleay, W. S. 1838. Illustrations of the Zoology of South Africa ; Annulosa. 75 pp., 4 pis. 

Milne-Edwards, H. 1837. Histoire Naturelle des Crustaces 2. 532 pp. Paris. 
Ortmann, a. E. 1896. Das System der Decapoden-Krebse. Zool. Jb., Jena, 9 : 409-453. 
Rathbun, M. J. 1904-5. Les Crabes d'Eau douce. Archs Mus. natn. Hist. nat. Paris (4) 

6 : 224-312 ; 7 : 159-321 ,* 18 pis. 
Williams, T. R. 1968. The taxonomy of the East African River-Crabs and their association 

with the Simulium neavei complex. Trans. R. Soc. trop. Med. Hyg., London, 62 : 29-34. 

S. F. Morris 

Department of Palaeontology 

British Museum (Natural History) 

Cromwell Road 

London SW7 5BD 

Accepted for publication 9 February 1976 

Fig. I 
Fig. 3 
Fig. 4 
Fig. 5 
Fig. 6 

Potamonautes (Lirrangopotamonautes) tugenensis sp. nov. 

Member C, Ngorora Formation, Miocene ; Kamwina, Kenya 
Holotype, Kenya National Museum. Dorsal carapace, xi-5. 
Paratype, Kenya National Museum. Sternum and ischium, x 3. 
Paratype, Kenya National Museum, x 1-2. 

Paratype, Kenya National Museum. Mesogastric and urogastric areas, x 3. 
Paratype, Kenya National Museum. Sternal 'knob', x 3. 

Member D, Ngorora Formation, Miocene ; Kalimale, Kenya 

Fig. 2. Kenya National Museum. Merus and carpus of left cheliped showing single carpal 
spine. X 1-75. 

Bull. Br. Mils. nat. Hist. (Geol.) 27, 4 


-, I 



:/'■ ■ 

%-rz L^ 




Serial sections confirm the occurrence of Cydothyris densleonis in the Upper Cretaceous of 
North America ; the known distribution of the genus is extended to Cahfornia. 

Mesozoic brachiopods have not been extensively described within the 
North American continent, and it is not surprising that Httle interest has 
been shown by palaeontologists in the few species of Rhynchonellidae and 
Terebratulidae already established in the literature. If, on the other hand, 
some of these records are further investigated, they can provide useful informa- 
tion regarding the ubiquitous nature of some genera and species of little-known 

Among early records of brachiopods from beds of known Cretaceous age are those 
of Whiteaves (1876- 1903) and Anderson (1902, 1958) who described rhynchonelloid 
brachiopod species which they referred broadly to the genus Rhynchonella. More 
recently Imlay (1937) described a species from the Lower Cretaceous of Mexico 
which he referred to Cydothyris ? subtrigonalis. This was the first citation, albeit 
tentative, of the genus Cydothyris from the American continent, but the accompany- 
ing transverse serial sections of the species do not support its assignment to this 

It was not until 1955 that Cooper, in a description of some brachiopods from the 
Cretaceous of Arizona, first accurately recorded the genus Cydothyris from North 
America, describing and figuring a species as C. americana from the Mural Limestone 
(Middle Albian) of the Bisbee Quadrangle. Until this positive record, the genus 
was considered to have been confined to the Cretaceous of western Europe 
and England where it is represented by species from the Aptian to Upper 

Anderson (1902) described and figured a rhynchonellid as Rhynchonella densleonis 
from the Middle Cretaceous of Shasta County (Horsetown Group), northern 
California. Later, in a description of Cretaceous beds of the Pacific coast (Anderson 
1958), he considered that beds within the Horsetown Group yielding R. densleonis 
were no younger than Lower Cenomanian in age and that their probable range was 
from Upper Albian to Lower Cenomanian. A specimen of R. densleonis Anderson 
has recently been obtained from the type section and is described here. From the 
internal structures seen in transverse serial sections of a specimen (Fig. 4) , it can be 
positively assigned to Cydothyris. 

Bull. Br. Mus. nat. Hist. (Geol.) 27, 4 : 301-304 Issued 28 October 1976 



Fig. 4. A series of fifteen transverse serial sections through the umbo of a dupHcate 
specimen of CyclothyHs densleonis Anderson from the Horsetown Group (Albian-Ceno- 
manian), Shasta County, Cahfornia. Geological Survey of America Locality No. 1051. 
The encircling deltidial plates around the foramen, the comparatively short dental lamellae 
and general morphology of the hinge-plates are typical of the genus. The numerals denote 
the distance in millimetres between each section, x 3. 



Subfamily CYGLOTHYRIDINAE Makridin 1955 

Genus CYCLOTHYRIS M'Coy 1844 

CyclothyHs densleonis (Anderson) 

Fig. 4 ; PI. 5, figs 1-6 

1902 Rhynchonella densleonis Anderson 
1958 Rhynchonella densleonis Anderson 

Description. External characters. Acutely biconvex Cyclothyris, 17-2 mm long, 
20 mm wide, 11-9 mm thick. Broadly triangular in general outline. The brachial 

72 ; pi. 7, figs 157, 158. 
Anderson : 86 ; pi. i, figs 3, 


valve is slightly inflated ; the pedicle valve has a broad, shallow sulcus originating 
from about the mid-line and widening anteriorly with a fairly extensive trapezoidal 
linguiform extension. Shell ornament consists of numerous radiating fine costae 
or costellae which are affected by a series of marked marginal plications creating a 
distinctive ornament ; similar ornament is characteristic of Cyclothyris antidichotoma 
(Buvignier) from the Upper Aptian and Lower Albian of France and England. The 
umbo is slightly produced and the sharp beak suberect. Distinct beak-ridges 
border an extensive interarea with exposed deltidial plates encircling a medium- 
sized foramen. 

Internal characters. The series of transverse serial sections given here (Fig. 4) 
for C. densleonis have been compared to those of Cyclothyris latissima (J. de C. 
Sowerby), the type-species from Faringdon, Berkshire, as figured by Owen (1962 : 46). 
The two series of sections appear almost identical in every detail. The character- 
istic deltidial plates encircling the foramen are shown to perfection, as well as the 
comparatively short dental lamellae in the pedicle valve. The shape, length and 
angle of deflection of the hinge-plates also agree, as do the depth of insertion and 
general quadrate shape of the hinge-teeth. 

Remarks. There is an unmistakable similarity between C. densleonis (Anderson) 
and C. antidichotoma (Buvignier), but it differs from the latter species in having finer 
and more numerous costellae, a more acutely triangular general outline, a narrower 
fold and sulcus, a more inflated brachial valve and considerably smaller overall 
dimensions. The marginal plicae in C. densleonis appear to be more marked within 
the sulcus and on the faint or incipient brachial fold, where there are usually three 
or four. 

In general outline it is more readily comparable to Cyclothyris mirahilis (Walker) 
from the Lower Albian of Leighton Buzzard, Bedfordshire, but differs in having 
more, but less acutely developed, marginal plicae, and a slightly more produced 
umbo ; it is less inflated than C. mirahilis. 

A species which shows a closer morphological affinity with the American form was 
described and figured by Panow (1969) as ? Cyclothyris antidichotoma (Buvignier), 
and occurs in Lower Cenomanian beds in the Cracow district of Poland. The 
specimen figured by Panow (1969 : pi. 109, fig. 9) has a wider anterior sulcus, a 
slightly less extensive linguiform extension, a less produced umbo and a less acutely 
inflated brachial valve. 

The importance of the present record is that it confirms that of Cooper (1955) in 
establishing Cyclothyris within the Albian and Cenomanian of North America and 
extends the distribution of that genus to include California. It may be possible in 
time to investigate the records of other workers, such as Whiteaves (1876- 1903), so 
that a clearer view of the distribution of Cyclothyris and similar brachiopod genera 
may be obtained. 


Anderson, F. M. 1902. Cretaceous deposits of the Pacific coast. Proc. Calif. Acad. Sci., 
San Francisco, (3) 2 i : 1-129, 12 pis. 




Anderson, F. M. 1958. Upper Cretaceous of the Pacific coast. Mem. geol. Soc. Am., New- 
York, 71 : 1-378, 75 pis. 

Cooper, G. A. 1955. New Cretaceous Brachiopoda from Arizona. Smithson. misc. Collns, 
Washington, 131 4 : 1-18, 4 pis. 

Imlay, R. 1937. Lower Neocomian fossils from the Miquihuana region, Mexico. /. Paleont., 
Chicago, 11 7 : 552-574. ^4 pls. 

Owen, E. F. 1962. The brachiopod genus Cyclothyris. Bull. Br. Mus. nat. Hist. (Geol.), 
London, 7 2 : 39-63, 2 pis. 

Panow, E. 1969. Contribution to the knowledge of the brachiopods from the Upper Cre- 
taceous of the Krakow district. Roczn. pol. Tow. geol., Krakow, 39 4 : 555-608, 4 pis. 

Whiteaves, J. E. 1876-1903. Mesozoic Fossils 1 : 1-415, 51 pis. Ottawa (Geol. Surv. 
Canada) . 

E. F. Owen, M.Phil.. M.I.Biol., F.L.S. 

Department of Palaeontology 
British Museum (Natural History) 
Cromwell Road 
London SW7 5BD 

Accepted for publication 9 February 1976 

Fig. la, b, c. 
BB 76200. 

Fig. 4a, b, c. 


Prefix BM(NH) - British Museum (Natural History), London 

MHNG - Museum d'Histoire Naturelle, Geneva, Switzerland 

Cyclothyris densleonis (Anderson) 
Cretaceous, Horsetown Group, Shasta County, northern California. BM(NH) 
X 2. 

Same locality and horizon. BM(NH) BB 76201. x 2. 

Cyclothyris antidichotoma (Buvignier) 
Lower Albian, Shenley Hill, Leighton Buzzard, Bedfordshire. BM(NH) 

Fig. 2a, b, c. 
BB 41495. X I. 

Fig. 2d. Enlargement of margin of shell of the same specimen as above, showing the typical 
antidichotoma ornament, x 2. 

Fig. 3a, b. Albian, Morteau, Doubs, France. Internal mould of a young specimen. 
CB 4746. X 2. 


ill. Br. Mus. nat. Hist. (Geol.) 27, 4 



















Cranial material is available for two mid-Tertiary flamingos, Phoenicopterus croiseti from the 
Oligocene and a new species from the Miocene, and this is compared with skulls of the three 
Recent genera. The skull of P. croiseti from the Upper Oligocene of France is partly recon- 
structed, using additional material. It is sufficiently distinct from the type species of Phoeni- 
copterus, the Recent P. ruber, to warrant its designation as the type species of a new genus, 
Gervaisia. Its bill is slender and less decurved than in Phoenicopterus, the mandibular sym- 
physis is shorter and the upper jaw is broad to the tip. New flamingo material from the 
Lower Miocene of Rusinga Island, Kenya, is referred to Phoenicopterus on the basis of palate 
structure. It is specifically distinct from P. ruber, the bill-shape being different and the size 
much smaller, and it is therefore described as a new species, P. aethiopicus. 


The Recent flamingos comprise four species separated into three genera, 
Phoenicopterus, Phoeniconaias and Phoenicoparrus, the last including both P. jamesi 
and P. andinus. Phoenicopterus ruber has three separate populations, P. r. 
antiquorum in Eurasia and Africa, P. r. ruber in the Caribbean and Central America 
and P. r. chilensis in southern South America. These are sometimes treated as 
separate species, but their bones differ only in size. All three forms have been used 
here for comparison but the skull of the last, Phoenicopterus ruber chilensis, has been 
chosen for the figures. 

Skulls representing the three genera were available for examination and are shown 
in lateral view (Fig. 5B-D). These are Phoenicopterus ruber, Phoeniconaias minor 
and Phoenicoparrus jamesi respectively. The three, in that order, show a progressive 
modification of the bill, probably linked with different feeding methods. In the 
first two we know of such differences, Phoenicopterus r^lber being a more generalized 
feeder, on minute Crustacea for the most part, and Phoeniconaias minor a more 
specialized feeder on blue-green algae. The bill-structure of Phoenicoparrus suggests 
a feeding method involving fine filtering similar to that of Phoeniconaias. 

The most generalized form, Phoenicopterus, has a longer and less angled bill. In 
the other two the lower mandible is stouter and shorter and is deflected downwards 
more vertically ; the bony structure is more laterally inflated and the mandibular 
symphysis longer. At the same time the upper jaw is laterally narrow (Fig. 6), and 
more sharply angled (Fig. 5), while the palatal ridge is a narrower and deeper wedge 
(Fig. 7). It is apparent that for the three Recent genera the palate and bill structure 
are diagnostic features, and cranial material of fossil species can be compared with 

Bull. Br. Mus. nat. Hist. (Geol.) 27, 4 : 305-314 Issued 28 October 1976 




Fig. 5. Lateral views of skulls of : A, Gervaisia croiseti (after Gervais) ; B, Phoenicopterus 
ruber chilensis ; C, Phoeniconaias minor ; D, Phoenicoparrus jamesi. The small arrows 
indicate the posterior margin of the mandibular symphysis ; the shaded portions of the 
upper mandible indicate the area of the palate visible laterally. 



Fig. 6. Anterior views of upper mandibles of : A, Gervaisia croiseti (after Gervais) ; 
B, Phoenicopterus ruber chilensis ; C, Phoeniconaias minor ; D, Phoenicoparrus jamesi. 

Fig. 7. Transverse section of palate, level with anterior end of nares, of 
pterus ruber ; B, Phoeniconaias minor ; C, Phoenicoparrus jamesi. 

A. Phoenico- 

these. Cranial material of fossil flamingos is available for only two forms, 
Phoenicopterus croiseti and a new form from Africa. 


Phoenicopterus croiseti Gervais 1852 
Figs 5A, 6A 

P. croiseti was described by Gervais from parts of two skulls and a tarsometatarsus 
from Clermont-Ferrand and Gergovie, Puy-de-D6me, France. The deposits at 
these localities are now thought by Thenius (1959) to be Upper Oligocene and not 
Lower Miocene as originally stated. More recently a specimen in the collection 
of the British Museum (Natural History) has been prepared ; it is no. A 2665 {ex 


2yd>2y), from the Croiset Collection purchased in 1848, and though said to be collected 
at Perignat, France, it is inscribed 'Gergovie'. The preparation has exposed most 
of the anterior part of a lower mandible and a separate small portion of the posterior 
end of the mandible in the region of the coronoid process. The mandibular 
sjrmphysis has undergone some lateral crushing. In shape, size and age the specimen 
is referable to the present species. With the help of it, using Gervais's plates but 
not his original specimens, it has been possible to reconstruct part of the skull of 
P. croiseti (Fig. 5A). 

The cranial region is too incomplete to be useful, but the bill shows distinctive 
characters (Figs 5 A, 6A). It is longer, narrower and less abruptly decurved than the 
bills of Recent species. The BM(NH) specimen appears deeper towards the tip of 
the lower mandible, but the broad tip of the upper jaw, present in Gervais's specimen, 
may cause the rami of the lower mandible to diverge more sharply dorsally and 
appear shallower in lateral view. As shown by the BM(NH) specimen the lower 
mandible is not only narrower and straighter than those of Recent species, with a 
shorter symphysis as indicated by the small arrows in Fig. 5, but the bony structure 
is also less inflated. That of Phoenicopterus shows distinct lateral inflation of the 
bone, especially in the deep mid-section of the mandible, and this inflation increases 
in Phoenicoparrus and Phoeniconaias . The posterior fragment of lower mandible 
(A 2665), which includes the coronoid process, has been crushed and provides little 
information other than that the process appears to have been weU developed and 
that the foramen a little below the process was present. 

The palatal ridge of P. croiseti is shallow and the upper jaw is down-curved but 
not angled. The end of the upper jaw differs from those of other flamingos in being 
broad right to the tip, those of the series of three Recent species listed above showing 
increased lateral constriction (Fig. 6) . 

From the shape of the bill P. croiseti would appear to have had a more generalized 
method of feeding than living flamingos. It differs more from the three extant 
genera than they differ between themselves. If the others are separated into different 
genera then croiseti requires a genus of its own. Apart from the cranial material we 
have little information on it ; the limb bones referred to this species by Gervais 
(1852), Milne-Edwards (1867-71) and Lydekker (1891) are similar in size and 
structure to those of Phoenicopterus ruber and are not outstandingly diagnostic. 
The following new genus is proposed. 

GERV AISI A gen. noY. 

Etymology. The genus is named after Paul Gervais who first described the 
species. It is feminine in gender. 

Type species. Phoenicopterus croiseti Gervais 1852 : 233-234 ; pi. 2, figs 4-5. 

Diagnosis. Bill long and slightly decurved but not angled. Palatal ridge 
shallow and apparently absent from anterior third of upper jaw. Upper jaw broad 
and dorsoventraUy flattened towards the tip, tapering only gradually. Similar in 
size to Phoenicopterus ruber. 



Occurrence. Transitional Oligocene/Miocene deposits of France and Germany 
(Brodkorb 1963 : 272). 


The collection of the British Museum (Natural History) contains some fragmentary 
avian material from the Lower Miocene of Rusinga Island in the Kenyan waters of 
Lake Victoria. A number of these fragments, mostly ends of long bones, appear 
referable to a single species of small flamingo, similar in size to the Recent 
Phoeniconaias minor. They consist of two mandibular fragments, one proximal and 
three distal ends of humeri, one distal end of a femur, three distal ends of tibiotarsi, 
and five proximal and two distal ends of tarsometatarsi. The limb bones, which are 
mostly broken or crushed to some degree, show Httle difference from those of 
Phoeniconaias minor save that an almost complete proximal end of a tarsometatarsus 
shows the calcaneal ridges of the hypotarsus to be similar in proximal view to those 
of Recent species but only half to two-thirds their length. 

The jaw fragments, however, differ more from Phoeniconaias minor in their 
structure. The larger fragment (Figs 8 A, 9 A) is the posterior part of the upper jaw 
with most of the narial apertures, embedded in matrix at its posterior end. Its 
most significant feature is the poorly-developed palatal projection with the double 
ridge along its centre. In this respect it resembles Phoenicopterus ruber so closely 
that it seems reasonable to include it in that genus (Fig. 9A-B). 

The other fragment is from the anterior end of the lower jaw. It is slender, 
slightly decurved, and heavily grooved in a manner more similar to that of 
Phoeniconaias. It tapers more gradually than does the bill of Phoenicopterus ruber 
in this region and may therefore have been longer and more slender ; it appears 
less deeply hollowed ventraUy towards the posterior and in comparison with P. ruber 
the bill may have been shallower in the region of the symphysis. The most marked 

Fig. 8. Dorsal views of basal area of upper mandibles of : A, Phoenicopterus aethiopicus 
(holotype) ; B, Phoenicopterus ruber chilensis ; C, Phoeniconaias minor ; D, Phoenico- 
parrus jamesi. 



Fig. 9. Lateral (right) view of posterior part of upper jaw and transverse section of palatal 
ridges of : A, Phoenicopterus aethiopicus ; B, P. ruber chilensis. 

Fig. 10. Transverse section of lower mandible, near tip, of : A, Phoenicopterus ruber 
chilensis ; B, Phoenicopterus aethiopicus (holotype) partly concealed by matrix. 

difference however is in the shape of the tomial edge of this region of the mandible. 
Recent species of flamingo, and as far as can be determined this applies also to 
Gervaisia croiseti, have a flattened dorsal edge to the tomium on which the flat edge 
of the upper jaw rests, although the latter is much reduced in Phoenicoparrus and 
Phoeniconaias. Phoenicopterus ruber has slender, incurving tomia on the lower jaw 
(Fig. loA). The new species has stout tomia slanting inwards on the inner surface, 
with a sharp dorsal ridge, and the outer surface bordering this ridge is lateraUy 
compressed to produce a shallow hollow (Fig. loB). The general appearance 
suggests that the anterior end of the upper jaw, at present unknown, may have 
overlapped the lower jaw. If this were so it would constitute a unique feature within 
the Phoenicopteridae and further taxonomic separation of this species might be 
justified. For the present, in view of the palatal similarity and the lack of confirma- 
tion of possible differences, we propose to treat this as a new species of the genus 
Phoenicopterus . 


Phoenicopterus aethiopicus sp. nov. 
Figs 8A, 9A, loB ; PI. 6 

Etymology. The species is named after the continent in which it occurs. 

Diagnosis. Small, slightly smaller than Phoeniconaias minor. Palatal ridge 
shallow and double, very similar to that of Phoenicopterus ruber. Tip of lower jaw 
narrow, slender and heavily grooved and with stout tomia compressed along the 
external side to produce a shallow groove, the appearance of which suggests that 
it might accommodate tomia of upper jaw. Tarsometatarsus with calcaneal ridge 
of hypotarsus short. 

Material. Holotype : the posterior part of an upper jaw, BM(NH) Pal. Dept. 
No. A 4382. Paratypes : BM(NH) Nos A 4383-4398, comprising the distal part of a 
lower jaw, two distal and five proximal ends of tarsometatarsi, three distal ends of 
tibiotarsi, one distal end of a femur, and three distal and one proximal end of humeri. 
Most material partly damaged or crushed. Collected and presented by the late 
Dr L. S. B. Leakey in 1935. 

Occurrence. Lower Miocene ; locality Rs 12, Rusinga Island, NE Lake 
Victoria, Kenya. 

Description. The holotype is the posterior portion of an upper jaw broken 
short proximal to the anterior end of the nares, and with some shattering of the 
internasal bridge. The posterior end terminates at the frontonasal hinge, where it 
is embedded in matrix. The bone is slender, widening dorsally at the posterior 
end. The culmen shows only slight curvature and is a little flattened where it 
terminates posteriorly. The nares are elongated and of fairly even width, rising 
slightly at the posterior end. The palate is flat posteriorly, with shallow paired 
ridges present along most of the length and increasing in depth anteriorly. 

The anterior end of the lower jaw (A 4384) is within the region of the symphysis 
and lacks the tip. It is slender, rounded and slightly decurved. Proximally it is 
flattened and slightly hollowed ventrally. Both ventral surface and sides have many 
deep longitudinal grooves. The internal surfaces are concealed by matrix, but the 
interior is deeply hollowed, with incurving tomia. The latter are thicker than those 
of P. ruber and project dorsaUy, the edges forming dorsal ridges. The sides of the 
jaw bordering the tomia are laterally compressed to form on each side a shallow 
hollow with a prominent lower margin, its appearance suggesting that the tomia 
of the upper jaw may have overlapped it. 

On the tarsometatarsus the hypotarsus is very short proximodistally, differing 
in this respect from those of Recent species and from the similar-sized P. minutus 
Howard 1955 of the Upper Pleistocene of Cahfornia. The distal end of the tarso- 
metatarsus is like that of Phoeniconaias, differing from that of P. novaehollandiae 
A. H. Miller 1963 of the Australian Miocene in the greater spread of trochleae and 
less clearly indicated articulation for digit i, from P.floridanus Brodkorb 1953 of the 
Florida Lower Pliocene in the proximally shorter trochlea for digit 2, and 
Phoeniconaias gracilis A. H. Miller 1963 of the AustraHan Early Pleistocene in the 
rounder and broader trochlea for digit 2 and more extensive articular surface on the 


trochlea for digit 4. The present species also differs from P. floridanus in having 
the distal end of the tibiotarsus narrower posteriorly, and from P. stocki L. Miller 
1944 of the Mexican Middle Pliocene in being proportionally narrower across the 
anterior intercondylar groove of this bone. The other known fossil flamingos, 
Phoeniconotius eyrensis A, H. Miller 1963 of the Australian Miocene and Phoenico- 
ptenis copei Shufeldt 1891 (see Shufeldt 1892) of the American Middle Pleistocene, are 
both considerably larger species. Except in the matter of size, however, the known 
differences between these various species are slight, and where critical cranial 
material is absent it is impossible to assess the true affinities adequately. 

Measurements. Holotype, A 4382 : dorsal length 27-2 mm ; ventral length 
28-1 mm ; depth at posterior end 17-0 mm, at posterior end of nares to tomium 
11-5 mm, to palate 14-3 mm, depth at anterior end to tomium 8-5 mm, to palate 
11-4 mm ; width of nares 5-7 mm ; width of posterior nasal strut 5-3 mm. 

Anterior portion of lower jaw, A 4384 : length 26-1 mm ; posterior width 12-4 mm, 
depth 9-8 mm ; width between tomia posteriorly 5-0 mm ; anterior width j'% mm, 
depth 7-5 mm ; width between tomia anteriorly 4-2 mm ; depth of lateral 
compression bordering tomia 2-2 mm. 

Proximal end of tarsometatarsus, A 4383 : width at proximal end 14-0 mm ; 
width of hypotarsus 7-9 mm ; anteroposterior depth of proximal end to hypotarsus 
15-8 mm ; depth of cotyla 8-6 mm ; proximodistal length of external calcaneal 
ridge 8-8 mm, of internal ridge 8-6 mm. 


We wish to thank Miss M. L. Holloway of the British Museum (Natural History) 
for preparing the line drawings. 


Brodkorb, p. 1953. A Pliocene flamingo from Florida. Chicago Acad. Sci. nat. Hist. Misc. 

124 : 1-4, 2 figs. 
1963. Catalogue of fossil birds, part i. Bull. Fla St. Mus. biol. Sci., Gainesville, 7 

4: 179-293- 
Gervais, p. 1848-52. Zoologie et Paleontologie Frangaises, 1. 271 pp. Paris. 
Howard, H. 1955. Fossil birds from Manix Lake, California. Prof. Pap. U.S. geol. Surv., 

Washington, 264-J : 199-206, pi. 50. 
Lydekker, R. 1 891. Catalogue of the fossil birds in the British Museum {Nat. Hist.). xxvii + 

368 pp., 75 figs. London. 
Miller, A. H. 1963. The fossil flamingos of AustraUa. Condor, Santa Clara, Calif., 65 : 289- 

299, 6 figs. 
Miller, L. 1944. A Pliocene flamingo from Mexico. Wilson Bull., Sioux City, Iowa, 56 : 

77-82, 2 figs. 
Milne-Edwards, A. 1867-71. Recherches anatomiques et paldontologiques pour servir a 

I'histoire des oiseaux fossiles de la France. 474 + 632 pp., atlas 200 pis (2 vols). Paris. 
Shufeldt, R. W. 1892. A study of the fossil avifauna of the Equus Beds of the Oregon 

desert. /. Acad. nat. Sci. Philad. (2) 9 : 389-425, pis 15-17. 
Thenius, E. 1959. Handbuch der stratigraphischen Geologic, 3, Tertiar. Teil 2, Wirbeltier- 

faunen. xi + 328 pp. Stuttgart. 



Dr C. J. O. Harrison, Ph.D. 

Suhdepartment of Ornithology 
British Museum (Natural History) 

C. A. Walker 

Department of Palaeontology 

British Museum (Natural History) 

Cromwell Road 

London SW7 5BD 

Accepted for publication 9 February 1976 


Phoenicopterus aethiopicus sp. nov. 

Holotype, BM(NH) A 4382. Views of the posterior portion of the upper jaw, x 2. A, right 
lateral. B, left lateral. C, dorsal. D, palatal. 

Paratype, BM(NH) A 4383. Views of the proximal end of a right tarsometatarsus, x 2. 
E, external. F, internal. G, proximal. 

Bull. Br. Mus. nat. Hist. (Geol.) 27, 





A new species of fossil pelican, Pelecanus aethiopicus , based on a tarsometatarsus and a scapula 
(both incomplete), is described from the Middle Pleistocene of Olduvai, Tanzania. It is com- 
pared with Recent and fossil forms. 

In the collection of the Department of Palaeontology of the British Museum 
(Natural History) are several fragments of bird bones collected from the Middle 
Pleistocene deposits at Olduvai, Tanzania. All the specimens are of birds associated 
with an aquatic environment, and are referable to Recent genera, but some may be 
sufficiently different from present-day forms to make it necessary to consider them as 
separate species. One such is the proximal part of a tarsometatarsus of a pelican 
which might have been a forerunner of one or more Recent species within the genus 


Suborder PELECANI Sharpe 

Family PELEGANIDAE Vigors 

Genus PELECANUS Linnaeus 

Pelecanus aethiopicus sp. nov. 

Plate 7 

Diagnosis. Proximal end of tarsometatarsus having internal edge of anterior 
surface with posterior curvature and with smooth, not abrupt slope to lip of internal 
cotyla. On internal anterior edge, the ridge marking proximal edge of ligamental 
furrow curves posteriorly only a little distal to level of hypotarsus. Distal edge of 
posterior aperture of inner proximal foramen only just proximal to level of distal 
edge of hypotarsus. Hypotarsus posteriorly prominent on internal side, with sharp 
slope towards internal side. Internal calcaneal ridge stout, with head only slightly 
prominent distally, and with slight ridge towards internal edge. Triangular bridge, 
tapering distaUy, over second tendinal groove formed by heads of next two calcaneal 
ridges. Internally the edge of this triangular head begins only a little anterior to 
edge of head of internal calcaneal ridge, and slopes externally, with some anterior 
curvature. External tendinal groove accommodated laterally, on the external side 
of hypotarsus, rather than on posterior surface (Fig. ii). 

Material. Holotype, proximal end of a right tarsometatarsus, British 
Museum (Natural History) Palaeontology Dept. No. A 4291. Paratype, proximal 
end of a right scapula. No. A 4292 in the same collection. 

Bull. Br. Mus. nat. Hist. (Geol.) 27, 4 : 315-320 Issued 28 October 1976 



CL 6 


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Pi r 

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6 P. 

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Locality and Horizon. Bed I, site FLKS, Olduvai Series (Middle Pleistocene), 
Olduvai Gorge, Tanzania (Leakey 1965 : 102). Collected and presented by the late 
Dr L. S. B. Leakey, 1935. 

Description. The holotype is the proximal end of a right tarsometatarsus 
(Fig. iiC) in good condition, but with the tendinal canals and anterior fossa filled 
with matrix and some crushing of the shaft distally. The external surface is broad 
and flat, terminating posteriorly in a distinct ridge, and anteriorly forming a more 
rounded ridge bordering the anterior fossa. The anterior surface has a distinct 
median metatarsal groove deepening towards the occluded fossa. At the distal edge 
of the fossa there is a small tibialis anticus tubercle on the inner side and a smaller 
ancillary pit next to it on the external side. 

The internal edge is more smoothly rounded than the external edge and shows 
some posterior curvature distally. At the proximal end it widens to accommodate 
a broad, short hollow just below the lip of the cotyla, presumably for the attachment 
of the proximal ligament ; the inner edge of this projects inwards across the proximal 
edge of the fossa. The more distal part of the internal edge is smoothed away where 
a ligamental band would have crossed it, this surface terminating proximally in a 
small ridge a little below the level of the tibialis anticus scar and curving posteriorly 
across the internal side (Fig. iiC). 

The internal surface is slightly curved and narrow proximally, to widen again 
just below the lip of the internal cotyla, which is broken in the present specimen. 
The proximal surface is roughly rectangular. The intercotylar prominence is 
elevated and bluntly rounded. Viewed anteriorly, it shows some external 
deflection ; viewed laterally it shows slight anterior prominence, continuing the line 
of the inner edge of the proximal ligamental attachment hollow, and dorsally slopes 
gradually towards the hypotarsus. The internal cotyla is narrower laterally and 
concave with a slight anterior tilt. The external cotyla is narrow and slopes 
anteriorly, its anterior edge curving down towards the anterior fossa. There is a 
narrow slanting intercotyla depression at the posterior inner edge of the internal 
cotyla, and a narrow, shallow groove crosses the posterior edge of this cotyla and 
extends to the opening of the second tendinal canal. 

The hypotarsus is large and stout. The main structure is the internal calcaneal 
ridge which is thick and prominent, although extending less far proximodistally 
than those of some Pelecanus species. Its posterior surface is flattened, projecting a 
little at the edges and sloping externally with a slight longitudinal ridge towards the 
internal side. Distally, below this overhanging surface the calcaneal ridge extends 
distally, decreasing in height. The internal surface of the ridge is irregular with small 
ridges and foramina. The posterior aperture of the inner proximal foramen, at the 
base of the inner calcaneal ridge just below the projecting surface, is large and 
irregular in shape. The anterior surface of the inner calcaneal ridge is concave, 
with a narrow projecting external edge. 

On the external side of the large inner calcaneal ridge there are two tendinal 
canals, one above the other, only one showing between the edge of the posterior 
surface of the ridge and the edge of the surface roofing the next tendinal canal. The 
second lateral tendinal canal, between the middle and outer calcaneal ridges, is 




roofed over. The middle calcaneal ridge is much shorter and slimmer than the 
inner, and only half as long. The roof of these forms a triangular-shaped surface, 
tapering distointernally and with a hollow above the tendinal canal, and has an 
external slant. The posterior opening of the outer proximal foramen is just distal 
and external to the posterior opening of the second tendinal groove. On the external 
side of the external calcaneal ridge there is a shallow lateral groove, a second groove 
diverging anteriorly from the distal end of this and passing across the proximal 
posterior corner of the external surface. Both these latter grooves are small and 
poorly defined, with a slight ridge between them which is eroded in the specimen. 

The paratype is the proximal end of a right scapula with furcular articulation 
broken off short, at a level with the eroded coracoid articulation. The shaft shows 
some crushing. The coracoid articulation is a prominent rounded facet projecting 
beyond the line of the anterior edge. The glenoid facet, which slants proximo- 
internally, is oval and elongate, with a small prominent lip along its outer edge, and a 
concavity in the bone on the dorsal surface above it. 

Measurements (in millimetres), (a). Holotype : proximal end of a right 
tarsometarsus, BM(NH) No. A 4291. 

Maximum length as preserved 53-9 

Maximum width across cotyla 24*5 

Minimum measurable width 16*2 

Depth of hypotarsus 29-5 

Depth at internal cotyla 15-5 

External depth of distal end to central posterior ridge 19-3 

Internal depth of distal end to central posterior ridge 15-6 

Depth of external side at level of tibialis anticus scar 13-4 

Depth of internal side at level of tibialis anticus scar 9-9 

Length of cap of internal calcaneal ridge 20-7 

Maximum width of cap of internal calcaneal ridge ii-2 

Length of second calcaneal ridge i6-o 

(b) Paratype : proximal end of a right scapula, BM(NH) No. A 4292. 

Maximum length of scapula as preserved 45-1 

Maximum width 26-2 

Width to internal edge of coracoid articulation 24-0 

Width of coracoid articulation 10-2 

Maximum thickness of shaft 5-9 

Width of shaft 11-3 

Width of glenoid facet 8-6 

Length of glenoid facet 14-0 

Comparison and comments. The following Recent specimens were available for 
examination. Pelecanus crispus 5, P. rufescens i, P. conspicillatus 2, P. onocrotalus 8, 
P. erythrorhynchus i, P. occidentalis 5. Of these, crispus (Fig. iiA), rufescens 
(Fig. iiB) and conspicillatus differ from aethiopicus in having on the external side of 
the large internal calcaneal ridge a series of three tendinal grooves of which the outer 
two are open, and in having a more posteriorly prominent calcaneal ridge. P. 
erythrorhynchus differs from aethiopicus in having the three tendinal grooves and a 
proximodistally long calcaneal ridge like that of onocrotalus but less abrupt distally. 
Its internal calcaneal ridge is, however, stout and only projects a short distance. 


P. onocrotalus differs from aethiopicus in its stouter, less projecting, and proximo- 
distally longer internal calcaneal ridge, and in its broader and less projecting 
hypotarsus. P. occidentalis is peculiar in that two discrete populations have different 
hypotarsi. The North and Central American population differs from aethiopicus in 
having the hypotarsus narrow and prominent, with two open tendinal canals 
posterior to a closed one, the more anterior of the two being partly enclosed. The 
southern population, P. 0. thagus of Chile and Peru, is a larger form in which the 
hypotarsus is much stouter and projects less. The external edge of the internal 
calcaneal ridge is extended externally, projecting beyond the tendinal canals. The 
two open posterior canals become displaced anteriorly and the more anterior of the 
two is completely enclosed. The general effect is as though the hypotarsus had been 
compressed onto the tarsometatarsal shaft with accompanying distortion. In view 
of this divergence in what is otherwise a specifically constant character there may be 
grounds for re-examining all characters of the two forms of occidentalis to determine 
whether their earlier separation as two species may be preferable. Both forms 
differ from aethiopicus in the characters mentioned. 

Hypotarsal material is available for only a few fossil pelicans. P. gracilis Milne- 
Edwards 1867, P. intermedius Fraas 1870 and P. odessanus Widhalm 1886 (listed by 
Lambrecht 1933 : 279) all have the narrower and more projecting hypotarsus typical 
of the crispus group ; on the last two the two open posterior tendinal grooves are 
apparent. The first two of these species are from the Miocene, the last from the 
Lower Pliocene. Of the three Pleistocene species already known P. halieus Wetmore 
1933 from North America, described from a radius, is very small and osteologically 
similar to erythrorhynchus , the Recent North American species. P. grandiceps 
De Vis 1906 and P. proavus De Vis 1892 (p. 444), found in Australia and described as 
similar to conspicillatus, the only Recent species occurring in that region, were 
described from various bones among which the proximal ends of tarsometatarsi 
were not represented. Present information suggests aethiopicus differs from all 
of these. 

Acknowledgements. We would like to thank Miss M. L. Holloway for making 
the line drawings, and Mr C. Keates for taking the photographs. 


De Vis, C. W. 1892. Residue of the extinct birds of Queensland as yet detected. Proc. Linn. 

Soc. N.S.W., Sydney, (2) 6 : 437-456 ; pis 23-24. 
1906. A contribution to the knowledge of the extinct avifauna of Australia. Ann. Qd 

Mus., Brisbane, 6 : 1-25, pis 1-9. 
Fraas, O. 1870. Die Fauna von Steinheim. Jh. Ver. vaterl. Naturk. Wurtt., Stuttgart, 

26 : 281-283 ; pi. 13, figs 3a-c, 4. 
Lambrecht, K. 1933. HandbuchderPalaeornithologie. xix+ 1022 pp., 4 pis, 209 figs. Berlin. 
Leakey, L. S. B. 1965. Olduvai Gorge jg5i-ig6i, 1. xiv+ii8 pp., 97 pis, 2 figs, 1 map. 

Milne-Edwards, A. 1867. Recherches anatomiques et paUontologiques pour servir a I'histoire 

des oiseaux fossiles de la France, 1 : 250-254 ; atlas 1 : pis 38-39. Paris. 
Wetmore, A. 1933. PHocene bird remains from Idaho. Smithson. misc. Collns, Washington, 

87, 20 : I -12, 8 figs. 



WiDHALM, J. 1886. Die Fossilen Vogel-knochen der Odessaer Steppen-Kalk Stein-briiche 
der Neuen Slobodka bei Odessa. Zap. novoross. Obshch. Estest., Odessa, 10, Beilage : 
4-6 ; pi. 5- 

Dr C. J. O. Harrison, Ph.D. 

Subdepartment of Ornithology 
British Museum (Natural History) 

C. A. Walker 

Department of Palaeontology 
British Museum (Natural History) 
Cromwell Road 
London SW7 5BD 

Accepted for publication 9 February 1976 

A, internal ; B, anterior 

Pelecanus aethiopicus sp. nov. 

Views of the proximal part of a right tarsometatarsus, BM(NH) No. A 4291. 
C, external ; D, posterior ; E, proximal, x i J. 

Bull. Br. Mus. nat. Hist. (Geol.) 27, 4 





A trilobite assemblage from the Mila Formation in northern Iran contains Chelidonocephalus 
preannulatus sp. nov., Hadragnostus edax sp. nov., Koldiniella mitella Sivov, Peronopsis fallax 
aff. minor (Brogger), Dorypyge sp., Parakoldinia ? sp. and Tsinania ? sp. It is of earhest 
Upper Cambrian age. The type species of Chelidonocephalus King (C. alifrons) and Iranoleesia 
King (7. pisiformis) are redescribed ; Iranoleesia falconi (King) is transferred to Anomocarella. 
Some problems in the classification of ptychoparioid trilobites are discussed. 


In NORTHERN Iran Cambrian rocks crop out in the east-west trending Alborz 
Mountain belt south of the Caspian Sea. In western Iran further Cambrian outcrops 
are known along a belt parallel to the Zagros Thrust north-west of Shiraz (Fig. 12). 
Fossils from the latter area have been described by King (1937), and more recently 
Kushan (1973) described the faunas and zones of the Mila Formation (Stocklin 
et al. 1964 : 20) in the Alborz Mountains. 

While surveying the Qazvin quadrangle north-west of Tehran (AnneUs et al. 1975), 
Dr R. G. Davies collected fossiliferous samples from within c. 10 m of the local base 
of the Mila Formation near Sanghabad (field locality RD 569), at lat. 36° 06' 43" N, 
long. 50° 37' 37" E. These samples yielded an assemblage of trilobites some- 
what ambiguous in age when compared with the faunas described by Kushan 
(1973). The assemblage includes well-preserved agnostids previously undescribed 
from Iran, and as a new species of Chelidonocephalus is present we have taken the 
opportunity to reassess Chelidonocephalus and associated genera and species known 
only from Iran, the affinities of which were obscure. 

The trilobites are preserved, with fragments of horny brachiopods, in a grey 
sparry limestone cut by many minor joints. Several, but not aU, show slight 
tectonic deformation. Although generally well preserved, the trilobites are all 
disarticulated and many were broken before fossilization. 

Reference is made to specimens in the Palaeontology Department, British 
Museum (Natural History) (It and In), the Geological Survey of India (GSI) and 
the Institute of Geological Sciences, London. 

We are grateful to Dr R. G. Davies for discussing the field occurrence of the 
fossils with us and to Mr S. F. Morris for technical assistance. A. W. A. Rushton 
publishes by permission of the Director, Institute of Geological Sciences. 

Bull. Br. Mus. nat. Hist. (Geol.) 27, 4 : 321-340 Issued 28 October 1976 


50 55 

Fig. 12. Sketch-map of part of Iran. Structural lines from Stocklin, 1968. Localities 
1-5 are from Kushan 1973 : i = Mila Kuh, 2 = Shahmirzad, 3 = Haanakdar, 4 = Ab- 
har, 5 = Qanli-Chapoghlu. Locality 6 = Sanghabad (R. G. Davies field Locality 569). 
Localities 7-10 are from King, 1937 • 7 = Chal-i-Sheh, 8 = Darreh Shu, 9 = Tang-i- 
Tehbud Bezuft, 10 = Ma'dan. Chelidonocephalus alifrons is recorded from localities 
I' 3' 4. 5. 7. 8, 9, Iranoleesia pisiformis from i, 3, 4, 5, 10 and Koldiniella mitella from 
3. 4. 5, 6. 


Kushan (1973 : 125) recognized the following zones in the Mila Formation. 

Tremadoc Saukia Zone 

C Albor sella Zone 

TT /- X. • J Kaolishania Zone 

Upper Cambrian < _ . . „ 

■^^ I Prochuangia Zone 

\Drepanura Zone 

Middle Cambrian 

j Dorypyge Zone 
1 Iranoleesia Zone 

Iranoleesia is confined to its zone and is accompanied by Anomocarella etc. 
According to Kushan the several Dorypyge species are confined to the Dorypyge 
Zone and some are associated with agnostids and Chelidonocephalus alifrons or C. sp. 
The Drepanura Zone is characterized by Drepanura [Spinopanura) or by Koldiniella, 


or both, and also contains C. alifrons, C. sp. and agnostids. Kushan did not allow 
for an overlap of the ranges of Dorypyge and Koldiniella, such as occurs in the 
present assemblage. 

We refer the present fauna to the Drepanura Zone because there are several 
specimens of Koldiniella mitella in the collection, and the presence of this species, 
of agnostids and of a species of Chelidonocephalus are typical of this zone. There is, 
on the other hand, only one specimen of Dorypyge and that is a different species from 
any recorded by Kushan from the Dorypyge Zone. On a priori grounds Kushan 
made a good case for the Middle Cambrian age of the Dorypyge Zone and the 
Upper Cambrian age of the Drepanura Zone, but as the new record implies an Upper 
Cambrian age for a Dorypyge species (a genus hitherto restricted to the Middle 
Cambrian) we here examine the age of the fauna in relation to the Middle- 
Upper Cambrian boundary. We accept the boundary as being between the Lejopyge 
laevigata Zone and the Agnostus pisiformis Zone, as reviewed by Daily & Jago 
(1975). Chelidonocephalus is restricted geographically and is of no help in wider 
correlation, nor is the Tsinania ? sp. because it cannot be convincingly compared 
with other described forms. Peronopsis fallax aff. minor resembles a subspecies 
known from the late Middle Cambrian of Scandinavia and P. fallax cf, minor from 
the A. pisiformis Zone in the Nuneaton district, England (Taylor & Rushton 
1972 : 19). Hadragnostus edax sp. nov. is comparable with H. las Opik from the 
eretes and quasivespa Zones of the Mindyallan Stage in Queensland : Opik regarded 
these horizons as Upper Cambrian but Daily & Jago (1975 : 538) suggested that the 
eretes Zone and part of the quasivespa Zone should be correlated with the top of the 
laevigata Zone, thus making H. las a topmost Middle Cambrian species. A third 
species, Hadragnostus modestus (Lochman), occurs in the Crepicephalus Zone of 
Montana (Lochman 1944) in beds equivalent to part of the A. pisiformis Zone. 
Thus these agnostids do not give a precise age as they are related to forms occurring 
both above and below the Middle-Upper Cambrian boundary. 

The only species of the present fauna found beyond the Mila Formation is 
Koldiniella mitella Sivov, which occurs in the lowest Upper Cambrian at Salair, 
western Siberia. Rozova (1968 : table 4) correlated this occurrence with the 
Kulyumbe 'superhorizon' of north-eastern Siberia which is referable to the 
Pedinocephalina-Toxotis (?) Zone, and this in turn was correlated with a level 
above the base of the A. pisiformis Zone by Daily & Jago (1975 : 546). K. mitella 
seems therefore to indicate an Upper Cambrian age, and with it goes the rest of the 
fauna, including the Dorypyge sp. We furthermore conclude that Kushan's choice 
of the Middle-Upper Cambrian boundary in the Mila Formation was apposite. 


Classification Problems. One of the reasons palaeontologists find the mass of 
'ptychoparioid' trilobites difficult to classify is that some persist in forcing genera 
into higher taxonomic groups on the basis of a single character in defiance of other 
characters which contradict such an assignment. Blindness and effacement of 
external features are thought to have been polyphyletically derived. Yet the blind 


Conocorj^hidae even now have the company of such distinct genera as Hartshillia 
and Meneviella thrust upon them despite Lake's observations that this ignores the 
characters of their thoraces and pygidia (1940 : 277). Similarly, Kobayashi, who 
recognized the polyphyletic character of effaced trilobites (1935 : 303), grouped 
nine varied genera in the Tsinaniidae (i960 : 397) but he himself regarded five of 
these as doubtful, with which we agree. We would add a sixth, Koldinia. This 
leaves the family with three closely related genera - Tsinania, Didyites and perhaps 
the poorly known Didyella : if Kobayashi is correct in regarding all these as effaced 
asaphiscids is there any need for a family Tsinaniidae at all? 

We have considered the genera likely to be related to Chelidonocephalus, 
Iranoleesia and Anomocarella below without regard to the (apparently haphazard) 
arrangement of the families in which they have been included. This has not led to 
a satisfactory understanding of the families concerned, and we have therefore been 
forced to use the category 'family uncertain'. 

Family AGNOSTIDAE McCoy 1849 
Genus HADRAGNOSTUS Opik 1967 

Type species. Original designation, H. las Opik 1967. 

Discussion. Hadragnostus has a distinctive cephalon with a long glabella and 
short preglabellar field. The front of the cephalon is transverse and straight, or 
even slightly emarginate in H. las (Opik 1967 : pi. 58, fig. 7) and H. modestus 
(Lochman 1944 : pi. 5, fig. 10) ; the new species H. edax, described below, has a 
straight or convex anterior margin. 

The pygidium of Hadrognastus differs from that of Peronopsis in that it has a 
terminal node. Baltagnostus has a terminal node but the pygidial border between 
the posterolateral spines is widened medially (i.e. slightly crescent-shaped) or has a 
pygidial 'collar'. Opik (1967 : 103) described the articulating device of H. las as 
simple, but the holotype of H. edax sp. nov. has an agnostoid articulation (Opik 
1967 : 72) which supports reference of Hadragnostus to the Agnostinae. 

Hadragnostus edax sp. nov. 
PI. II, figs 3-15 

Derivation of name. Latin, edax, greedy, referring to the swollen pygidial axis. 

Holotype. A well-preserved pygidium. It 13463 (PI. 11, figs 7, 8, 12). 

Type locality and horizon. Mila Formation, Sanghabad, Taleghan Range, 
Alborz Mountains, northern Iran. 

Figured material. Cephala It 13468 (PI. 11, fig. 3), It 13483a (PI. 11, figs 9, 
10), It 13483b (PI. II, figs 14, 15), It 14023 (PI. II, fig. 11). Pygidia It 13463 
(holotype, PI. 11, figs 7, 8, 12), It 13471 (PI. 11, fig. 13), It 13472 (PI. 11, 
figs 4-6). 


Description. Cephalic outline tends to be subquadrate, length about four fifths 
of width. Glabella three quarters of cephalic length, bilobed ; anterior lobe 
relatively small, nearly twice as wide as long, narrower than posterior lobe and one 
third of its length or less. Posterior lobe parallel-sided, rounded or slightly pointed 
behind, without lateral furrows or a median node, but some specimens have a faint 
posterior node (PI. 11, fig. 15). Basal lobes triangular, inflated, connected behind 
glabella. Each cheek (excluding border) as wide as posterior glabellar lobe, tapering 
strongly forwards ; in front of the glabella the cheeks are narrower than the length 
of the anterior glabellar lobe and are faintly separated by a partial preglabellar 
median furrow which forms a shallow depression just in front of the glabella and 
shallows or fades out forwards. Laterally and frontally the cheeks curve down 
steeply to the border which is mainly flat and horizontal but has a narrow outer 
rim and is curved under at the edge. 

Thorax unknown. 

Pygidium strongly convex, transverse, length about three quarters of width. 
Axis large, extending back to overhang the posterior border furrow. At anterior 
end axis is just less than half the total pygidial width, and widens slightly backwards. 
The articulating furrow has a depression either side of the median line, giving a 
weakly diplagnostid character. A median tubercle at the highest point has its 
posterior end just in front of the mid-length of the axis. Irregularities in the axial 
furrow indicate the merest traces of lateral furrows opposite the front and back of 
the median tubercle. A small terminal node lies well above the border furrow. 
Each flank about two fifths as wide as axis in the larger specimens but proportionally 
wider - nearly half the axial width - in a small specimen about i-i mm long. Flanks 
slope steeply down to the border, narrow backwards, separated behind axis. Border 
broad and thick, of even width between the small posterolateral marginal spines. 
Surface smooth. 

Discussion. The present form is very similar to Hadragnostus las Opik 
(1967 : pi. 58, figs 6-10) but has a slightly shorter and narrower anterior glabellar 
lobe. In H. las this lobe is more than one third of the length of the posterior lobe 
but in the present form it is less than a third. H. las has a stronger and wider 
preglabellar median furrow. The pygidial flanks are narrower, less than half the 
width of the axis, whereas in H. las they are more than half, and in Opik's reconstruc- 
tion (1967 : 103, text-fig. 24) the posterolateral marginal spines are shown as larger 
than those of H. edax. 

One figured cephalon of Hadragnostus modestus (Lochman 1944 : pi. 5, fig. 10) is 
typical of the genus : the anterior glabellar lobe is narrower than the posterior lobe 
but it is longer than in H. edax. The preglabellar median furrow is narrower than 
that of H. las and stronger than in H. edax. The basal lobes of H. modestus are 
smaller than in either of the other species. Lochman's other figured cephalon has 
a more rounded outhne, the posterior glabellar lobe is more tapered and the 
preglabellar field is longer ; it may represent another genus. The holotype and 
paratype pygidia of H. modestus are meraspids with the median tubercle 
at the mid-length of the pygidium. Full comparison with H. edax cannot be 


Family QUADRAGNOSTIDAE Howell 1935 
Genus PERONOPSIS Hawle & Corda 1847 

Type species. By monotypy, Battus integer Beyrich 1845. 

Peronopsis fallax (Linnarsson 1869) aff. minor (Brogger 1878) 
PL 12, figs 1-14 

Figured material. Cephala It i348oa-b (PI. 12, figs i, 2), It 13480c (PL 12, 
figs 3, 4), It i348od (PL 12, fig. 14), It 14024 (PL 12, figs 9, 11), It 14026 (PL 12, 
fig. 10). Pygidia It 13467 (PL 12, fig. 6), It 13469 (PL 12, figs 12, 13), It 13485 
(PL 12, figs 5, 8), It 14025 (PL 12, fig. 7). Some twenty unfigured cephala and 
pygidia also present. 

Description. Cephalon rounded in front, nearly as long as wide. Glabella 
about 0*7 of cephalic length, bilobed ; anterior lobe rounded in front, slightly 
narrower than, and less than two fifths the length of, the posterior lobe. Trans- 
glabellar furrow narrow at external surface, broader in exfoliated specimens. 
Posterior glabellar lobe has faint lateral furrows at two fifths of its length from the 
front and a faint median node, generally elongate, at or in front of its mid-length ; 
bluntly rounded behind. Basal lobes triangular but with a forward extension in 
some specimens which indents the sides of the glabella, somewhat as in Ptychagnostus 
atavus (Tullberg) ; this feature is not seen in exfoliated specimens. Cheeks about 
as wide as glabella, narrowing forwards slightly, confluent in front. A pit 
resembling an incipient preglabellar median furrow is seen in front of the glabella in 
some specimens. Cheeks slope steeply down to border in small or exfoliated 
specimens, less steeply in the largest specimens. Border furrow is a curve where 
the cheeks flatten out to form a border, and is broad in large specimens. Border is 
a narrow convex rim. 

Thorax unknown. 

Pygidium convex, subquadrate, length about 0-9 of width. Axis about two fifths 
of total width, parallel-sided with a tendency to narrow slightly at the anterior 
third. Posterior end generally bluntly pointed but may be rounded, especially in 
large specimens, not reaching border. Median tubercle at anterior two fifths ; 
traces of lateral furrows opposite anterior and posterior ends of tubercle. Articu- 
lating device simple, basic (Opik 1967 : 72). Flanks (pleural lobes excluding border) 
about half the width of axis but proportionally narrower in specimens more than 
2 mm long, and always confluent behind axis although the connection is narrow in 
some specimens. Border furrow broad and shallow. Border broad, of even width 
between the strong posterolateral marginal spines. Surface smooth. 

Discussion. In most features the present form falls within the wide range shown 
by Westergard's (1946 : pi. 3, figs 3-7) figures of P. fallax minor. It differs from 
many Peronopsis species in that the pygidial flanks are confluent behind the axis, 
and resembles P. fallax minor but not P. fallax ferox (Tullberg) in the character of 
the cephalic border (Westergard 1946 : 38). The cephala agree with Westergard's 
figures although the glabella is generally longer than in his fig. 4 and the anterior 


glabellar lobe is proportionally longer than in his fig. 3 ; neither figure shows a 
preglabellar depression or elongated Piychagnostus-like basal lobes. A specimen of 
P. fallax of. minor (PL 12, fig. 15) from the A. pisiformis Zone of central England 
(Taylor & Rushton 1972 : 19) has a weak preglabellar depression but differs in having 
a longer anterior glabellar lobe and smaller basal lobes ; the pygidial axis is evenly 
rounded behind. Probably compression has caused the lateral margins of the 
pygidium to converge backwards slightly. 

Some of the pygidia of 'Agnoshts' simplexiformis Rozova (1964 : 24) resemble that 
of the present form. Rozova figured no cephala of '^ .' simplexiformis but Lazarenko 
& Nikiforov (1968 : pi. 3, figs 1-4) showed that both cephalon and pygidium have 
constricted acrolobes (Opik 1967 : 68). Judging from these figures we would transfer 
'A.' simplexiformis to Agnostoglossa Opik (1967 : 145). 

Family DORYPYGIDAE Kobayashi 1935 
Genus DORYPYGE Dames 1883 
Type species. Original designation, Dorypyge richthofeni Dames 1883. 
Discussion. About forty specific names have been applied to Dorypyge, sensu 
strido, but according to Kobayashi's revision (i960 : 347) many of these are 
synonyms. Kushan (1973) has discussed six forms from Iran. 

Dorypyge sp. 

PI. II, figs I, 2 

Material. One fragmentary pygidium about 7 mm long, It 13461. 

Description. Outline, excluding spines, trapezoidal, width nearly 1-5 times 
length. Axis only slightly tapered backwards, composed of three well-defined 
rings, a fourth less well defined, and a rounded bulbous terminal part. Flanks 
slightly narrower than axis anteriorly, marked by three pairs of narrow pleural 
furrows but no interpleural grooves. Border broad, the anterior pleural furrows 
crossing it indistinctly. Medially, the border forms a narrow transverse connection 
behind the axis. Convex parts of surface covered with sparse coarse granules. The 
marginal spines are distinctive although not all are preserved. The pygidium 
follows the common plan of Dorypyge species in having four pairs of main marginal 
spines anterior to the largest pair, but the present form is unusual, and perhaps 
unique, in showing a small subsidiary spine anterior to each of the first three of the 
'main' spines ; their bases make the outline of the pygidium jagged. Whereas the 
thickness of the 'main' spines increases from the first to fourth pair, the subsidiary 
spines are progressively thinner from the first to third pair. The fifth pair of 
marginal spines is represented only on the left by a scar where the spine is broken 
off ; it was twice as thick as the fourth spine. The outline of the posterior part of 
the pygidium suggests the former presence of a substantial sixth pair of spines. The 
doublure, which is partly exposed, is a smooth band parallel to the margin and 
convex ventrally. 


Discussion. The present form differs from all species so far described in having 
subsidiary pygidial spines. However Dr P. Jell has told us of certain Kootenia 
species which also show 'subsidiary' marginal spines which point more nearly 
ventrally and are correspondingly difficult to observe. Their presence suggests 
that similar spines may be found in other Dorypygidae. In other respects the 
present form resembles the subspecies D. richthofeni laiwuensis Kobayashi, which 
lacks interpleural grooves, and the form figured by Sun (1924 : pi. 2, figs 3 c, d) 
which Kobayashi (i960 : 348) referred to D. r. laevis. Sun's fig. 3d shows that the 
marginal spines increase in size backwards and also shows sparse granulation, but 
it differs in having slightly wider flanks and only three axial rings. The present 
specimen is distinguished from D. iranensis Kushan and D. khademi khademi Kushan 
by the granular surface, and from D. khademi papulosa Kushan by the absence of 
interpleural grooves. The fragment illustrated by Kushan (1973 : pi. 28, fig. 8) as 
D. richthofeni n. subsp. aff. richthofeni differs slightly in having wider axial ring 
furrows and pleural grooves, and a wider (sag.) post-axial part of the border. 

Family ASAPHISGIDAE Raymond 1924 

Genus IRANOLEESIA King 1955 

{Pro Irania King 1937, ^^w De Filippi 1863) 

Diagnosis. Asaphiscid trilobites with glabella slightly tapering, truncate 
anteriorly. Posterior two pairs of glabellar furrows deep, inner half of iP bifurcate, 
posterior branch directed obliquely backwards. Occipital ring may be subdivided 
with lateral lobes. Postocular fixed cheeks small (sag., trans.). 

Type species. Original designation, Irania pisiformis King 1937. 

Discussion, The type species (and, with the removal from Iranoleesia of Irania 
falconi King 1937, the only species) of Iranoleesia is redescribed below. Kobayashi 
(1967 : 439) indicated that Iranoleesia may be a junior synonym of Hundwarella 
Reed 1934, from the Cambrian of Kashmir. The two specimens on which the 
type and only species of Hundwarella is based, both cranidia, show apparently 
bifurcate inner ends of the iP glabellar furrows (Reed 1934 : pi. II, figs 5, 6) which 
connect across the middle of the glabella. This is not the case in Iranoleesia 
pisiformis, but the Kashmir specimens are slightly crushed, and the apparent 
transverse furrows may be the result of compressing bifurcate furrows of Iranoleesia 
type. A similar effect may be observed on flattened specimens of Hypermecaspis 
(compare Fortey 1974 : pi. 13, fig. i with Lake 1913 : pi. 7, fig. 3). Even if this is 
accepted Iranoleesia pisiformis differs from Hundwarella in having a transversely 
truncate front margin of the glabella, the preglabellar field only half as wide (sag.), 
and a strongly punctate surface sculpture. Since the significance of such characters 
in discriminating asaphiscid genera is still imperfectly appreciated, Iranoleesia is 
therefore retained here pending a revision of the whole group. For example, the 
genus Iniotoma Opik 1967, from the Mindyallan, has apparently similar glabellar 
furrows (Opik 1967 : 232, fig. 81) to those of Iranoleesia, but they are less impressed 
and the eye ridges are less distinct (1967 : pi. 11, figs 1-3). While it seems probable 


that these genera are separated more by geography than by morphology, a revision 
of the genera is beyond the scope of the present account. The genus Protohedinia 
Endo 1937 (the type species of which has been illustrated by Chang 1963 : pi. i, 
fig. 9) has glabellar furrows similar to those of Iranoleesia and has palpebral lobes 
similarly placed, eye ridges of like prominence, and border and preglabellar fields 
of similar proportions. It is difficult to see why it has been included in a separate 
family Tengfengiidae by Chang (1963 : 458). 

Iranoleesia pisiformis (King 1937) 
PI. 9, figs 6, 8-10, 12 
1937 lycit^ici' pisiformis King : 12-13 ; pi. II, figs 6a-c 

Diagnosis. Iranoleesia species with pitted surface sculpture. Anterolateral 
corners of glabella angulate. Anterior border furrow without plectrum. 

HoLOTYPE. Although not named as such, it is clear from the plate description 
and measurements given by King (1937 : 13 ; pi. II, fig. 6) that he intended the 
incomplete cranidium, GSI 16305, as holotype of this species. We figure a cast of 
this original on PI. 9, fig. 6. 

Type locality and horizon. Ma'dan, locaHty 12 of King (1937). The exact 
stratigraphic relationship of this species to the new Chelidonocephalus fauna is not 
known for certain but Kushan recorded /. pisiformis only from horizons below those 
with Koldiniella mitella, suggesting that the Chelidonocephalus fauna is slightly the 

Figured material. Cranidia In 36890 (cast of holotype, PI. 9, fig. 6), 
In 36910 (PI. 9, fig. 8), In 36911 (PI. 9, fig. 10), In 36912 (PI. 9, figs 9, 12). 

Description. Only cranidia known with certainty, and available material 
largely decorticated, but otherwise well preserved in a limestone matrix. General 
cranidial outline subquadrate, with anterior margin not greatly protruded, glabella 
but slightly elevated above fixed cheeks, sloping downwards anteriorly into pre- 
glabellar field. Glabella (with occipital ring) almost rectangular, extending to 
three quarters cranidial length, tapering very gently forwards, so that the axial 
furrows enclose an angle of about 15°. Anterolateral corners of the glabella are 
obtusely angulate, resulting in a truncate anterior glabellar margin. Four pairs of 
lateral glabellar furrows, of which the anterior two pairs are shallow and probably 
would not be visible on the dorsal surface of the cuticle. iP has its outer end 
opposite the midpoint of the palpebral lobe, its outer third running transversely or 
slightly posteriorly, at which point it bifurcates into a short, slightly anteriorly- 
directed branch, and a longer, strongly backward-directed posterior branch, the 
inner end of which terminates at about one third glabellar length. 2P, with its outer 
end almost opposite the front of the palpebral lobe, is arcuate, the inner end curving 
backwards in hue with the inner end of iP. 3P and 4P opposite the eye ridge, the 
former isolated within the glabella, transverse, the latter short, slightly forward- 
inclined, placed laterally to 3P. Occipital furrow deep, forward-curved medially, 


and shallowing laterally ; occipital ring with indistinctly defined lateral lobes and 
prominent median tubercle. Axial furrows distinct, as is the preglabellar furrow on 
the smaller cranidium (PI. 9, fig. 8), although on larger specimens this furrow becomes 
shallower to almost effaced medially. 

Intraocular fixed cheeks about half width (trans.) of adjacent glabella, horizontal, 
Preglabellar field about one sixth length of glabella (sag.), scarcely downsloping 
medially ; that part of the fixed cheeks in front of the eye ridge slopes downwards 
more steeply and is about twice the length (exsag.) of the preglabellar field (seen in 
dorsal view). The eye ridges are strong on internal moulds, converging forwards 
at about 65° to the sagittal line. Palpebral lobes of about same width as the eye 
ridges, in a posterior position such that the transverse line connecting their anterior 
limits crosses the glabella at two thirds its length ; they are of length (exsag.) one 
third, or slightly less, that of glabella. Anterior border furrow defining change in 
slope from preglabellar field to broad, backward-sloping anterior border, the width 
of which (sag., exsag.) is similar to that of the preglabellar field. Postocular cheeks 
poorly shown by present material. 

Genus ANOMOCARELLA Walcott 1905 

Diagnosis. Asaphiscid trilobites with glabellar furrows faint or absent ; 
preglabellar field less than one fifth glabellar length ; anterior border furrow with 
narrow (trans.) plectrum ; anterior border flat. Pygidium with distinct flat 

Type species. Original designation, Anomocarella chinensis Walcott 1905. 

Discussion. The species described below as Anomocarella falconi w3iS originally 
assigned to Irania by King (1937), and hence subsequently to Iranoleesia. 
Kobayashi (1967 : 493) preferred to regard the species as belonging to Grandioculus 
Cossmann, The type species of that genus, G. megalurus (Dames) (Dames 1883 : 20 ; 
pi. I, figs 7, 8, 10, Walcott 1913 : 192 ; pi, 18, figs 9a-f) has large, somewhat 
posteriorly placed palpebral lobes, weakly defined glabellar furrows and a flat 
border like that of the Iranian specimens, but the border furrow lacks a plectrum, 
which is also the case in Iranoleesia pisiformis. The type species of Anomocarella 
is A. chinensis Walcott 1905, the lectotype of which was carefully identified by 
Endo & Resser (1937 : 164-165). This cranidium is only one of the specimens used 
by Walcott (1913 : pi. 20, figs 3, 3a-d, 4, 4a) to illustrate A . chinensis, and all the others 
are from a different locality from the lectotype (pi. 20, fig. 3c) and include more than 
one species (Endo & Resser 1937 : 165), It is the other material (the specimens 
of Walcott 1913 : pi. 20, figs 3, 3e) that is used to illustrate the type species of 
Anomocarella in the Treatise (Howell in Harrington et al. 1959 : 292). Endo & 
Resser emphasized the presence of a plectrum as 'the most distinctive generic 
feature' of Anomocarella, with which we agree. The major point of difference 
between the Iranian material and the type species is the shorter preglabellar field 
of the latter ; this is not regarded as of generic importance, as it is a variable feature 
in other species assigned to Anomocarella by Endo & Resser (1937), Further back- 
ward migration of the plectrum (or further shortening of the preglabellar field) 


results in the broadly backward-deflected border furrows seen in Mapania (Opik 
I96i:fig. 53). 

Anomocarella and allied genera are sometimes referred to a separate family 
Anomocarellidae Hupe 1955. Since the characters pertinent to the division of 
genera within the Asaphiscidae and AnomocareUa-grow-^ are unclear at best, and 
there is no phyletic concept on which to base separate families, Anomocarella is here 
retained doubtfully within the Asaphiscidae. 

Anomocarella falconi (King 1937) 
PI. 9, fig. II ? ; PI. 10, figs 1-5, 8 
1937 I yania falconi King : 13-14 ; pi. II, figs ya-f. 

Diagnosis. An Anomocarella species with transverse width of plectrum half to 
two thirds that of anterior margin of glabella ; preglabellar field of moderate width ; 
dorsal surface of cuticle minutely granulose. 

HoLOTYPE. An external mould of a cranidium, a cast from which was figured 
by King (1937 : pi. II, fig. 7a), GSI 16306. This cast is here refigured on 
PL 10, fig. I. 

Type locality and horizon. Mila Formation at Darreh Shu (lowest beds), 
locality 8 of King (1937), whence Chelidonocephalus alifrons is also recorded (King 
1937 • 17)- 

Figured material. Cranidia In 36891 (cast of holotype, PL 10, fig. i). In 36908 
(PL 10, fig. 3), In 36909a (PL 10, fig. 2), In 36909b (PL 10, figs 4, 5, 8). Doubtfully 
assigned pygidium In 36892 (cast, PL 9, fig. 11). 

Description. Cranidium of low convexity, downward-sloping in front of eye 
ridges, glabella not greatly vaulted above intraocular cheeks. Maximum cranidial 
width at posterior margin about one and a half times transverse width at anterior 
border. Glabella extends to about three quarters cranidial length, and tapers 
uniformly forwards, the axial furrows enclosing an angle between 15° and 20°. 
Anterolateral corners of glabella rounded ; axial furrow deeper than preglabellar 
furrow, and both better defined on internal moulds. Occipital and glabellar 
furrows faint, best seen as smooth, slightly depressed areas on specimen retaining 
exoskeleton (PL 10, fig. 8). iP to 3P almost touch axial furrows ; distance (exsag.) 
between iP and 2P equal to that between iP and the occipital furrow, but less 
than that between 2P and 3P ; 3P is level with the point at which the eye ridges 
touch the glabella, transverse ; iP and 2P slope slightly backwards. 4P is very 
close to 3P but external to it, continuing in line with the furrow defining the posterior 
of the eye ridge. Occipital ring widest (sag.) medially, defined by broad, shallow 
furrow, of which only the transverse, median section is visible on the internal mould 
(PL 10, fig. 3) ; outer ends of this section deepened into internal muscle scars. 
External surface shows narrow parafrontal band in front of midpart of frontal 
glabellar lobe ; this band extends further laterally on the internal mould, but neither 
on dorsal nor on ventral surfaces is the connection with the eye ridge displayed. 



Palpebral lobes of half length of glabella, anterior limit opposite outer ends of 3P 
furrows ; palpebral rims broad, defined by shallow palpebral furrows which are 
outward-bowed medially. Maximum transverse width of intraocular cheek half 
width of adjacent glabella. Eye ridges not visible abaxially on dorsal surface of 
exoskeleton, but their confluence with the frontal lobe of the glabella is clearly 
shown by anterior shallowing of the axial furrows. The eye ridges, where seen, are 
only slightly oblique, virtually transverse on the smallest cranidium. Postocular 
fixed cheeks narrow, width (exsag.) less than that of occipital ring, bisected by 
strong border furrow. Narrow posterior border carries small articulating socket (?) 
at about mid-width, maximum transverse width of border less than that of occipital 
ring. Preglabellar field of length (sag.) between 0-14 and 0-17 times glabellar length 
on available material. Anterior border furrow marks an abrupt change in slope 
anterolaterally, where it is gently bowed forwards ; medially there is a distinct 
plectrum, of transverse width about half to two thirds that of frontal lobe of glabella. 
Laterally the wide anterior border is horizontal to slightly declined, but in front of 
the plectrum more nearly carries forwards the downward slope of the preglabellar 
field. There are faint indications of caeca crossing the border furrow. The dorsal 
surface of the exoskeleton is covered with minute granules. 

An incomplete pygidium (PI. 9, fig. 11) was assigned to this species by King 
(1937 : pi. II, fig. 7b). It has a convex rhachis showing three or four axial rings and 
a broad flattened border. It also resembles the posterior part of the pygidium 
here tentatively assigned to Chelidonocephalus preannulatus sp. nov. (p. 338), and 
it must be admitted as a possibility that the pygidium is that of C. alifrons. Our 
material is inadequate to resolve the question. 

Discussion. Anomocarella falconi differs from the type species A. chinensis 
(Walcott 1913 : pi. 20, fig. 3c. Endo & Resser 1937 : pi. 34, fig. 6) in having less 
oblique eye ridges (due to an anterior forward limit of the palpebral lobe) and a 
longer (sag., exsag.) preglabellar field; the surface sculpture of A. chinensis is 
described as punctate. Of the many species assigned by Walcott (1913 ; some 
re-iUustrated in Lu et al. 1965 : pis 59-61) to Anomocarella only the type species has a 
plectrum, and the others should be excluded from the genus. One of the species 
from the Mapan Formation, A. concava Endo & Resser (1937 : 167 ; pi. 35, fig. 8), 
approaches A . falconi closely in the proportions of preglabellar field and plectrum. 
It differs only in having a glabella that hardly tapers forwards, and anterior limits 
of the palpebral lobes that approach the glabella more closely. 

Genus KOLDINIELLA Sivov 1955 

Type species. Original designation, K. mitella Sivov 1955. 

Discussion. This genus is characterized by a broad, flat cephalic border and 
effacement of the other cephalic characters. Traces of the axial furrow suggest that 
the glabella extends to or nearly to the anterior border (Kushan 1973 : pi. 31, fig. 3) ; 
this feature, the more backward eyes and the transverse pygidium distinguish 
Koldiniella from the more effaced forms of Maryvillia and Bloimtia (Rasetti 1965 : 


pis 9, 10). Koldiniella is similar to but more effaced than the asaphiscid Peishania 
Resser & Endo from the late Middle Cambrian ; it may even have been derived from 
a form like P. luhrica Chang (1957 : pi. i, fig. 2) which has a comparable frontal 
border but has the axial furrows and occipital furrow more distinct. Peishania 
(and Liopeishania Chang 1963, to which P. luhrica was transferred in Lu et al. 1965) 
have longer pygidia than Koldiniella mitella. The dorsal surface of the cranidium 
of Liopeishania spannensis Palmer & Gatehouse is more effaced than other species 
and approaches Koldiniella in this respect. It differs in having a shorter anterior 
border which is curled down and under in front (Palmer & Gatehouse 1972 : pi. 4, 
fig. 10). 

Koldiniella has been variously grouped with the families lUaenidae, lUaenuridae 
and Tsinaniidae on account of its effaced cephalon, but there are no characters 
uniting it convincingly with any of these families and the frontal border is a feature 
distinguishing it from all (and from Kingstonia). It perhaps comes closest to the 
lUaenuridae because Illaenurus has a transverse pygidium and in Macelloura the 
glabella reaches to the frontal border. 

Koldiniella mitella Sivov 1955 

PL 10, figs 6, 7, 9, 10, 15, 13 ?, 14 ?, 16 ? 

Figured material. Cranidium It 14021 (PI. 10, figs 6, 7) ; pygidia It 13477 

(PI. 10, figs 9, 10), It 14022 (PI. 10, fig. 15) ; hypostome assigned to the species 

It 13506 (PL 10, fig. 16) ; cranidium doubtfully assigned, It 13476 (PL 10, figs 13, 14). 

Ten unfigured specimens also present. 

Discussion. Kushan (1973 : 149) redescribed this species using material from 
Iran. Material in the present collection is identical. A hypostome tentatively 
referred to K. mitella (PL 10, fig. 16) has a gently convex median body divided by a 
faint transverse furrow. The lateral border widens out posterolaterally but is 
narrow posteromedially. The surface is smooth except posterolaterally where it is 
striated parallel to the margin. This hypostome cannot be that of Dorypyge 
because it differs from that referred to Dorypyge aenigma (Westergard 1948 : pi. 2, 
fig. 7) ; the smooth surface and striated margin recall Koldiniella mitella rather than 
the associated Chelidonocephalus preannulatus. 

K. mitella is distinguished from K. convexa Lazarenko by the longer and flatter 
anterior border and by the weaker axial furrows at the base of the glabella (Rozova 
1964 :pL 15, figs 1-5). 

K. prolixa Lazarenko {in Lazarenko & Nikiforov 1968 : pi. i, figs 16-19) has a 
longer, more salient frontal border and a much longer pygidium, nearly as long as 

One small cranidium 17 mm long (PL 10, figs 13, 14) has a much narrower (sag.) 
frontal border, less than a tenth of the length of the cranidium. It may possibly be a 
juvenile form oiK. mitella because among larger specimens the border seems to have 
grown more rapidly than the rest of the cranidium, but the border is convex instead 
of being flat or even slightly concave as it is in specimens of K. mitella 3 mm or more 
in length. The small cranidium may therefore be referable to Parakoldinia Rozova 


i960. It is most closely comparable with P. plana Rozova (i960 : pi. 7, figs 1-5), a 
broad and not especially convex species with a distinct frontal border, horizontal in 
front view, which does not show strong striae. Unfortunately the present specimen 
is too different in size for an exact comparison to be made, but Rozova's fig. 3 shows 
the border furrow more sharply cut and the border dropping abruptly downwards in 
front, whereas in the present specimen the border curves down more gradually in 

Genus TSINANIA Walcott 1914 
Type species. Original designation, Illaenurus canens Walcott 1905. 

Discussion. Kobayashi (1935 : 303) has suggested that Tsinania is an effaced 
member of the Asaphiscidae. 

Tsinania ? sp. 

PI. 10, figs II, 12, 17-19 

Figured material. Cranidia It 13493 (PL 10, figs 11, 12), It 13458 (PI. 10, 
figs 17-19)- 

Description. Cranidium convex. Axial furrows effaced at external surface but 
a trace of their ends seen on the exfoliated specimen. The base of the glabella is a 
little more than half as wide as the cranidium. No trace of occipital or glabellar 
furrows. In palpebral view the middle of the eyes Hes just behind the mid-length. 
Palpebral furrow very faint. Preocular sutures diverge forwards slightly but curve 
inwards across the anterior border. Postocular sutures diverge backwards obhquely, 
straight. A very faint border furrow separates off a narrow, barely differentiated 
frontal border. Pleuroccipital furrow weak. Surface smooth. 

Discussion. These specimens agree closely with Tsinania canens (Walcott 1913 : 
pi. 23, fig. 2) except that the eyes seem farther forward, the base of the glabella is 
wider and there is a very faintly marked border. Kobayashi (1952) revised Tsinania 
canens and showed that the glabella is less than half as wide as the cranidium. No 
published figures show a frontal border. Tsinania ? sp. also resembles Pletho- 
meto(fus obtusus Rasetti (Longacre 1970 : pi. 6, figs 14, 15) except that the latter has 
a well-marked occipital furrow and curved postocular sutures. Plethometopus 
convergens (Raymond) has only a weak occipital furrow but the preocular sutures 
converge forwards (Longacre 1970 : pi. 3, figs 11, 12). Plethopeltoides lepidus 
Lazarenko {in Lazarenko & Nikiforov 1968 : pi. 9, figs 1-7) has a better-marked 
occipital furrow and is less convex. Wanwanoglohus convexus (Kobayashi 1966 : 265 
and text-fig. 4) has a much more convex cranidium. We know of no Cambrian 
trilobite which matches the present cranidium in all details. 


Diagnosis. Cranidium opisthoparian, 'ptychoparioid', with glabellar furrows 
(other than occipital) not incised. Broad false border furrow delimiting flattened 


or gently convex anterior cranidial border. True border furrow crosses preocular 
cheek, cut by preocular suture at sutural mid-length (exsag.), inner ends of border 
furrow curve backwards to define the plectrum. Narrow (sag.), transverse inflated 
band in front of glabella. Broad palpebral lobes between one third and half 
glabellar length, posterior limits opposite outer parts of occipital furrow defining 
narrow (exsag.) postocular cheek. Width of intraocular cheek less than that of 
adjacent glabella. Eye ridges subdued. 

Type species. Original designation, C. alifrons King 1937. 

Discussion. Chelidonocephalus was placed in the Alokistocaridae in the Treatise 
(Howell in Harrington et at. 1959 : 238), but other possibilities must now be con- 
sidered. Chang (1959 : 223) allied Chelidonocephalus with a number of Middle 
Cambrian genera from China, including Poshania Chang 1957 (type species P. 
poshanensis Chang 1957), Peichishania Chang 1957 (type species Eymekops rect- 
angularis Endo & Resser 1937), Inouyella (type species I. peiensis Endo & Resser 
1937), Ordosia (type species 0. fimbricauda Lu 1954) and Taitzuia (type species 
T. insueta Endo & Resser 1937). All these genera share with Chelidonocephalus the 
backward-curved border furrow (termed below the 'true' border furrow), although 
the development of the border itself is variable. In the Treatise (Harrington et al. 
1959) Chelidonocephalus, Ordosia, Inouyella and Taitzuia are assigned to four different 
superfamilies, while Chang (1959 : 223) indicates that they may be included within 
a single family Namanoiidae Lermontova 1951, based on the Lower Cambrian 
genus Namanoia from eastern Siberia, which has a similar disposition of furrows 
on the cranidial margin. We believe that the form of the border furrows is assuredly 
significant at generic level, but that its significance at family level is more contentious. 
The backward-curving border furrows are present, for example, on Llanoaspis 
walcotti Resser (Rasetti 1965 : pi. 8, fig. 14) from the Upper Cambrian of Tennessee, 
which is certainly unrelated to Chelidonocephalus and the other genera listed above, 
having distinctively different facial sutures. Similarly Namanoia, with its quadrate 
glabella and lack of parafrontal band, is unlikely to be closely related to Chelidono- 
cephalus, which therefore cannot be assigned to the Namanoiidae. 

We consider that Chelidonocephalus is most nearly related to Poshania, which is 
identical in the combination of true border furrow and parafrontal band (Chang 
1959 : pi. 2, figs 4, 5), and differs significantly only in lacking a distinct false border 
furrow. The resemblance is perhaps more apparent when comparing small cranidia 
of Chelidonocephalus (PI. 9, fig. 5) with adult Poshania as the whole border region 
is then of similar proportions. Peichishania, Inouyella, Ordosia and Taitzuia are 
probably unrelated to these two genera ; Peichishania was assigned by Opik (1967 : 
221) to the Auritamidae, Ordosia by Lu (1954) to the Leiostegiidae (and Taitzuia 
closely resembles Ordosia). Inouyella, with its tapering glabella with narrow, 
backward-directed furrows (Endo & Resser 1937 : pi. 46, figs 9, 14), is probably 
unrelated to any of the foregoing. 

The Alokistocaridae have been included within the concept of the Papyriaspididae 
by Opik (1961 : 149). He (Opik 1967 : 298) includes Poshania (misspelt Poshauia) 
in the family Mapaniidae of his superfamily Rhyssometopacea, which might therefore 


be expected to include Chelidonocephalus also. We briefly discuss below (p. 337) some 
reasons for treating a rhyssometopacean affinity with caution. 

This discussion shows to what an extent the familial (and superfamilial) concepts 
are capable of varied interpretations in these late Middle to early Upper Cambrian 
forms. For the time being, therefore, we have to leave Chelidonocephalus as of 
uncertain family. 

Chelidonocephalus alifrons King 1937 
PI. 8, figs 1-5 

Diagnosis. Chelidonocephalus species with punctate surface sculpture. Anterior 
border of cranidium gently convex. Palpebral lobes farther from glabella, and 
convexity lower than that of C. preannulatus nov. sp. (p. 338). 

Lectotype. King (1937) did not specify a holotype for this species, but of his 
original syntypic series that figured in his pi. II, fig. 8 A corresponds with the measure- 
ments cited in the text, and was probably intended as the type. We therefore 
select this specimen, GSI 16311, as lectotype. 

Type locality and horizon. The lectotype is associated with Anomocarella 
falconi (King) from a limestone 2300 feet (700 m) below the local top of the Cambrian, 
at Chal-i-Sheh (loc. 3 of King 1937 : text-fig. i). The evidence from Kushan's 
(1973) faunal lists with Chelidonocephalus is that a late Middle Cambrian or an 
earliest Upper Cambrian age is most probable for these beds. 

Figured material. Cranidia In 36893 (cast of lectotype, PL 8, fig. i), In 36895 
(PL 8, figs 2, 3, 5), In 36896 (PL 8, fig. 4). 

Description. Cranidia only identified with certainty, these being well pre- 
served in limestone. Convexity is low, the whole surface of the cranidium sloping 
gently downwards and forwards from the occipital ring. The largest specimen, a 
fragmentary cranidium, probably had a cephalic length of about 20 mm. Maximum 
transverse width, across posterior margin, about one and a half times the width 
across the anterior cranidial margin. Glabella tapering gently forwards, axial 
furrows enclosing an angle of about 30°, extending to about 0-7 of the cranidial 
length (sag.) ; front margin truncate. Glabellar furrows absent, but muscle in- 
sertion areas are represented by smooth areas lacking the pitted sculpture which 
covers the rest of the glabella. There is a pair of large smooth areas centred on the 
lateral parts of the occipital furrow, and three (? four) pairs of ill-defined smooth 
areas on the flanks of the glabella. The occipital furrow is shallow except where 
deepened into pits near its lateral extremities. Postocular and preocular fixed 
cheeks downsloping ; intraocular cheek horizontal. Palpebral lobes about one 
third glabellar length, with posterior limits slightly in front of points at which 
the occipital furrow meets the axial furrows. Transverse width of the intraocular 
cheek at anterior hmit of palpebral lobe is about 07 of the width of glabella at same 
section. Palpebral furrows are faint even on internal moulds. Eye ridges are 


faint on the external surface of the cuticle, but clearly visible on internal moulds 
(PL 8, fig. 4) ; their adaxial limits indent the course of the axial furrows. Posterior 
border furrow deep and transverse ; convex posterior border of width (exsag.) sub- 
equal to that of furrow. In front of the glabella there is a narrow (sag., exsag.) 
transverse inflated band, defined posteriorly by the preglabellar furrow and an- 
teriorly by the true anterior cranidial border furrow. This latter follows a trans- 
verse, gently curved course a little anterior to the front margin of the glabella, 
medially deflected sharply backwards when approaching the anterolateral corners 
of the glabella, to form the median plectrum (Opik 1967 : 58), this area being inflated 
above the genal areas adjacent to it. The anterior border furrow is cut by the 
anterior branch of the facial suture at, or slightly in front of, the anterior sutural 
mid-length. In front of the plectrum the preglabellar field continues a gently 
downward slope into a broad, shallow false border furrow. The gently convex 
anterior border widens medially, where it occupies one third to one quarter of the 
total preglabellar length. Internal moulds are strongly caecate on frontal part of 
cranidium ; caeca pass over both border furrows, but are not visible behind eye 
ridge. Dorsal surface of cranidium is finely pitted except in furrows and over 
muscle insertion areas. In front of the true border furrow, the pits are replaced by 
more densely crowded, minute granules. 

Discussion. The description of 'true' and 'false' border furrows used above 
needs clarification. It is contended that the posterior of the two transverse furrows 
crossing the preglabellar area represents the border furrow as understood in other 
trilobites, and that the anterior furrow in Chelidonocephalus is independently 
generated. Homologically, all that area anterior to the true border furrow should 
be termed 'anterior border', but for convenience the area is divided into preglabellar 
field, 'false' border furrow and anterior border in line with other trilobites. The 
identification of the posterior of the two furrows with the anterior border furrow of 
other trilobites is based on the fact that this furrow has a median plectrum, and 
that the plectrum is invariably developed on the anterior border furrow. Since 
plectra are developed in a number of groups which are, at least on present classi- 
fications, only distantly related, its presence may be taken as showing the 
position of the true anterior border furrow. The important defining character 
of Chelidonocephalus is the presence of two anterior furrows. It is interesting 
to note that some forms comparable in this respect are to be found in the 
superfamily Rhyssometopacea Opik (1967 : 272) described from the Mindyallan of 
Australia, for example Rhyssometopus princeps Opik (1967 : pi. 27, fig. 1-3 ; text- 
fig. 96), if we can accept Opik's 'frontal wrinkle' {ibid. : text-fig. 93) as 
homologous to our true border furrow. However, rhyssometopaceans show impor- 
tant differences from Chelidonocephalus, notably in having large palpebral lobes 
close to the glabella, and we are unable to classify our species with the Australian 

A comparative description of Chelidonocephalus preannulatus sp. nov. is given 
below, which extends the concept of the genus to include forms with larger palpebral 
lobes, closer to the glabella. They do not, however, approach rhyssometopaceans 
in this respect. 


Chelidonocephalus preannulatus sp. nov. 
PI. 8, figs 6-8 ; PI. 9, figs i, 2, 5. 3 ?, 4 ?, 7 ? 

Diagnosis. Chelidonocephalus species with granulate surface sculpture. An- 
terior border broad and flat. Palpebral lobes larger, placed closer to the glabella 
than in C. alifrons, and convexity and furrow incision generally greater. 

Derivation of name. Latin - 'with a ring in front' - referring to the inflated, 
transverse preglabellar band. 

HoLOTYPE. BM(NH) It 13503, incomplete cranidium partly preserving exo- 
skeleton, figured on PI. 9, fig. 2. 

Type locality and horizon. Mila Formation, Sanghabad, Taleghan Range, 
Alborz Mountains, northern Iran. 

Figured material. Cranidia It 13478 (PI. 8, figs 6-8), It 13504 (PL 9, fig 5) ; 
free cheek It 13499 (P^- 9- %• i) » hypostoma, tentatively assigned. It 14020 (PI. 9, 
figs 3, 4) ; pygidium, tentatively assigned, It 13460 (PI. 9, fig. 7). Six unfigured 
fragments also present. 

Discussion. Since this species is similar to the type species C. alifrons in most 
proportional characters, a detailed description need not be repeated here. C. pre- 
annulatus differs from the type species in the following characters. 

(i) Convexity (sag., exsag.) is greater, this being due to the increase in the down- 
ward deflexion of the fixed cheek in front of the palpebral lobes. 

(ii) Anterior border is flat, not gently convex, forming a broad, horizontal rim 
around the cranidial margin. The border occupies about half (sag.) the preglabellar 
width, although the false border furrow, except on small cranidia, is ill defined. 

(iii) Anterior branches of the facial sutures appear more divergent in dorsal view, 
this being due to the greater downward inclination of the preocular cheeks. 

(iv) Length of palpebral lobes about half length of glabella. They are also closer 
to the glabella, such that the transverse width of the intraocular cheek at the anterior 
limit of the palpebral lobe is about half width of glabella at same section (0-7 times 
in C. alifrons). 

(v) Furrows more deeply incised. 

(vi) Surface sculpture of fine granules on glabella. Smooth areas on the glabella 
of one specimen (PI. 8, figs 6-8) display the muscle insertion areas rather better 
than C. alifrons. Lateral occipital impression deep, producing a lateral narrowing 
of the occipital ring. iP subcircular, indistinct, isolated within glabella near mid- 
line ; 2P of similar size and form adjacent to axial furrow opposite mid-part of 
palpebral lobe ; 3P opposite anterior end of palpebral lobe, narrow and transverse ; 
4P of similar form, slightly anteriorly directed. 

A free cheek (PI. 9, fig. i) shows a prominent eye socle, faint caeca radiate from 
the eye and there is a lateral continuation of the true and false border furrows 
which describe a ridge on the border. The counterpart shows the base of a strong 
genal spine. 

A small hypostoma (PI. 9, figs 3, 4) may belong to C. preannulatus. It is 
different from that assigned to Koldiniella, and from that of DoryPyge, so that 


Chelidonocephalus is the most likely polymerid known from the present fauna to 
which it could belong. A distinctive feature is a prominent anterior hump on 
the middle body. 

An incomplete associated pygidium is tentatively referred to this species because it 
shows traces of similar granular sculpture. The axis is wider than the flanks, tapers 
back, is rounded behind but tends to pass into a postaxial ridge ; it is composed of 
4-5 rings and a terminal piece. Flanks show two distinct and one less clear pleural 
furrows. No interpleural furrows. There is a suggestion of a flattened lateral border. 

Annells, R. N., Arthurton, R. S., Bazley, R. A. B., & Davies, R. G. 1976. Explanatory 

text of the Qazvin and Rasht Quadrangles Map. Geological Quadrangles Nos E3 and E4. 

Geological Survey of Iran, Teheran. 
Chang W.-T. i957- Preliminary note on the Lower and Middle Cambrian stratigraphy of 

Poshan, Central Shantung. Acta palaeont. sin., Peking, 5 : 29-31, 2 pis. 

1959. New trilobites from the Middle Cambrian of North China. Acta palaeont. sin., 

Peking, 7 : 215-236, 4 pis. 

1963. A classification of the Lower and Middle Cambrian trilobites from North and 

Northeastern China, with description of new families and new genera. Acta palaeont. sin., 
Peking, 11 : 475-487, 2 pis. 

Daily, B. & Jagg, J. B. 1975. The trilobite Lejopyge Hawle and Corda and the Middle - 

Upper Cambrian boundary. Palaeontology, London, 18 : 527-550, pis 62-63. 
Dames, W. 1883. Cambrische Trilobiten von Laiu-Tung. In Richthofen, F. von, China 

4 : 3-33, Pis I, 2. Berlin. 
Endo, R. & Resser, C. E. 1937. The Sinian and Cambrian formations and fossils of southern 

Manchoukuo. Bull. Manchur. Sci. Mus., Mukden, 1. ix + 474 pp., 73 pis. 
FoRTEY, R. A. 1974. The Ordovician trilobites of Spitsbergen. I. Olenidae. Sky. norsk 

Polarinst., Oslo, 160. 81 pp., 24 pis. 
Harrington, H. J. et al. 1959. Systematic descriptions. Class Trilobita. In Moore, R. C. 

(ed.). Treatise on Invertebrate Palaeontology , O. xix + 560 pp., 415 figs. Lawrence, Kansas. 
King, W. B. R. 1937. Cambrian trilobites from Iran (Persia). Mem. geol. Surv. India 

Palaeont. Indica, Calcutta, 22 (5). 22 pp., 2 pis. 

1955- Iranoleesia, new trilobite name. Geol. Mag., Hertford, 92 : 86. 

KoBAYASHi, T. 1935. The Cambro-Ordovician formations and faunas of South Chosen. 
Palaeontology, part 3. /. Fac. Sci. Tokyo Univ. sec. 2, 4 : 49-344, pis 1-24. 

1952. On Mansuyia and the Tsinanidae. Trans. Proc. palaeont. Soc. Japan, Tokyo, 

N.s. 5 : 145-154. Pl- 13- 

i960. The Cambro-Ordovician formations and faunas of South Korea. Part VII, Palae- 
ontology VI. /. Fac. Sci. Tokyo Univ. sec. 2, 12 : 329-420, pis 19-21. 

1966-67. The Cambro-Ordovician formations and faunas of South Korea. Part X, 

Stratigraphy of the Chosen Group in Korea and South Manchuria and its relation to the 
Cambro-Ordovician formations of other areas. Section B. The Chosen Group of North 
Korea and Northeast China. /. Fac. Sci. Tokyo Univ. sec. 2, 16 : 209-311. Section C. 
The Cambrian of Eastern Asia and other parts of the Continent. Ibid. : 381-535. 

KusHAN, B. 1973. Stratigraphie und Trilobitenfauna in der Mila- Formation (Mittelkambrium- 
Tremadoc) im Alborz-Gebirge (N-Iran). Palaeontographica, Stuttgart, A 144 (4-6) : 113- 
165, pis 26-34. 

Lake, P. 1906-46. British Cambrian trilobites. Palaeontogr. Soc. {Monogr.), London. 350 
pp., 47 pis. 

Lazarenko, N. p. & NiKiFOROV, N. I. 1968. [Assemblages of trilobites from Upper Cambrian 
strata on the River Kulyumbe (northwestern Siberian Platform).] Uchen. Zap. nauchno- 
issled. Inst. geol. arkt., Leningrad, Pal. Biostrat. 23 : 20-80, pis 1-15. [In Russian.] 



Lermontova, E. V. 195 1. [Lower Cambrian trilobites and brachiopods from Eastern Siberia.'] 

222 pp., 21 pis. Moscow (Vses. nauchno.-issled. geol. Inst. (VSEGEI)). [In Russian.] 
LocHMAN, C. 1944- /w LocHMAN, C. & DuNCAN, D. Early Upper Cambrian faunas of Central 

Montana. Spec. Pap. geol. Soc. Am., Baltimore, 54. 181 pp., 19 pis. 
LoNGACRE, S. A. 1970. Trilobites of the Upper Cambrian Ptychaspid Biomere, Wilberns 

Formation, Central Texas. Rep. Invest. Bur. econ. Geol. Univ. Texas, Austin, 66 : 1-70. 
Lu Y.-H. 1954. Two new trilobite genera of the Kushan Formation. Acta palaeont. sin., 

Peking, 3 : 425-438, i pi. 

et al. 1965. [Fossils of each group of China: Chinese Trilobites.'] 766 pp., 134 pis, 

2 vols. Peking. [In Chinese.] 

Opik, a. a. 1961. The geology and palaeontology of the headwaters of the Burke River, 
Queensland. Bull. Bur. Miner. Resour. Geol. Geophys. Aust., Melbourne, 53. 249 pp., 
24 pis. 

1967. The Mindyallan fauna of north-western Queensland. Bull. Bur. Miner. Resour. 

Geol. Geophys. Aust., Canberra, 74. (i) 404 pp. ; (2) 167 pp., 67 pis. 

Palmer, A. R. & Gatehouse, C. G. 1972. Early and Middle Cambrian trilobites from Antarc- 
tica. Prof. Pap. U.S. geol. Surv., Washington, 456-D. 37 pp., 6 pis. 

Rasetti, F. 1965. Upper Cambrian trilobite faunas of northeastern Tennessee. Smithson. 
misc. Collns, Washington, 148 (3). 127 pp., 21 pis. 

Reed, F. R. C. 1934. Cambrian and Ordovician fossils from Kashmir. Mem. geol. Surv. 
India Palaeont. Indica, Calcutta, 21 (2). 38 pp., 2 pis. 

Resser, C. E. & Endo, R. 1937. ^^ Endo, R., & Resser, C. E., q.v. 

RozovA, A. V. i960. [Upper Cambrian trilobites of the Salayr.] Trudy Inst. Geol. Geofiz. 
sib. Otdel., Novosibirsk, 5. 116 pp., 8 pis. [In Russian.] 

1964. [Biostratigraphy and description of trilobites of the Middle and Upper Cambrian of 

the north-west Siberian Platform.] 148 pp., 19 pis. Moscow, Akad. Nauk SSSR, Sibir. 
otdel. Inst. Geol. Geofiz. ('Nauka'). [In Russian.] 

1968. [Biostratigraphy and trilobites of the Upper Cambrian and Lower Ordovician of 

the north-west Siberian Platform.] Trudy Inst. Geol. Geofiz. sib. Otdel., Moscow, 36. 
196 pp., 17 pis, 5 tab. [In Russian.] 

Sivov, A. G. 1955. In Khalfina, L. L. [Atlas of guide forms in the fossil faunas and floras 

of western Siberia.] 327 pp. Moscow. [In Russian.] {Fide Kushan, not seen.) 
Stocklin, J. 1968. Structural history and tectonics of Iran : a review. Bull. Am. Ass. 

Petrol. Geol., Tulsa, 52 : 1229- 1258. 
, RuTTNER, A. & Nabavi, M. 1964. New data on the Lower Paleozoic and Pre-Cambrian 

of North Iran. Rep. geol. Surv. Iran, Teheran, 1. 29 pp., i pi. 
Sun Y.-C. 1924. Contributions to the Cambrian faunas of North China. Palaeont. sin., 

Peking, B 1 (4). 109 pp., 5 pis. 
Taylor, K. & Rushton, A. W. A. 1972 [1971 on cover]. The pre-Westphalian geology of 

the Warwickshire Coalfield. . . . Bull. geol. Surv. Gt Br., London, 35. 152 pp., 12 pis. 
Walcott, C. D. 1905. Cambrian faunas of China. Proc. U.S. natn. Mus., Washington, 

26 : I -106. 
1913. The Cambrian faunas of China. In Willis, B. et al.. Research in China 3 : 1-276, 

pis 1-24. Carnegie Inst., Washington (publ. 54). 
Westergard, a. H. 1946. Agnostidea of the Middle Cambrian of Sweden. Sver. geol. 

Unders. Avh., Stockholm, C 477. 140 pp., 16 pis. 
■ 1948. Non-Agnostidean trilobites of the Middle Cambrian of Sweden. I. Sver. geol. 

Unders. Avh., Stockholm, C 498. 32 pp., 4 pis. 

R. A. FoRTEY, Ph.D. 

Department of Palaeontology 

British Museum (Natural History) 

Cromwell Road 

London SW7 5BD 

A. W. A. Rushton, B.A., Ph.D. 
Institute of Geological Sciences 
Exhibition Road 
London SW7 2DE 

Accepted for publication 9 February 1976 


Chelidonocephalus alifrons King 1937 (p. 336) 

Mila Formation, Chal-i-Slieh, Loc. 3 of King 1937 

Fig. I. Plaster cast of lectotype (selected herein), incomplete cranidium, x 4. Specimen 

figured by King 1937 • pl- 2, fig- 8a, GSI 16311 (cast In 36893). 

Figs 2, 3, 5. Cranidium retaining cuticle, dorsal and lateral views x 5 ; Fig. 5, detail of right 
side of glabella showing surface sculpture of fine pitting, x 12-5. In 36895. 

Fig. 4. Large exfoliated cranidium, showing caeca and form of facial suture, dorsal view, x 3. 
In 36896. 

Chelidonocephalus preannulatus sp. nov. (p. 338 ; see also PL 9, figs 1-5, 7) 
Mila Formation, near Sanghabad, Alborz Mountains 

Figs 6-8. Cranidium retaining cuticle, oblique lateral and dorsal views, x 4 ; Fig. 8, detail of 
right side of glabella to show sculptural difference from C. alifrons, x 12. It 13478. 

Bull. Br. Mus. nat. Hist. (Geol.) 27, 



Chelidonocephalus preannulatus sp. nov. (p. 338 ; see also PL 8, figs 6-8) 

Mila Formation, near Sanghabad, Alborz Mountains 

Fig. I. Fragmentary free cheek, x 8. It 13499. 

Fig. 2. Holotype, partially exfoliated cranidium, x 8. It 13503. 

Figs 3, 4. Hypostoma attributed to this species, ventral view, and lateral view to show 
anterior hump, x 10. It 14020. 

Fig. 5. Smallest cranidium, x 15. It 13504. 

Fig. 7. Pygidium probably attributable to this species, fragmentary, x 4. It 13460. 

Iranoleesia pisiformis (King 1937) (p. 329) 
Mila Formation, Ma'dan, Loc. 12 of King 1937 

Fig. 6. Cast of holotype, incomplete cranidium, x 5. Original specimen figured by King 
1937 : pi. 2, fig. 6a, GSI 16305 (cast In 36890). 

Fig. 8. Two cranidia, one retaining part of the cuticle and showing punctate surface sculpture, 
X 6. In 36910. 

Figs 9, 12. Internal mould of incomplete exfoliated cranidium, dorsal and lateral views, x 10. 
Note punctation on internal surface. In 36912. 

Fig. 10. Internal mould of cranidium, x 5. In 36911. 

Anomocarella falconi (King 1937) ? (p. 331) 
Mila Formation, Darreh Shu, I-oc. 8 of King 1937 

Fig. II. Cast of incomplete pygidium, x 6 ; original figured by King 1937 • P^- 2, fig. 7b, and 
there attributed to A. falconi, but possibly belonging to Chelidonocephalus alifrons (p. 332). 
GSI 16307 (cast In 36892). 

Bull. Br. Mils. nat. Hist. (Geol.) 27, 4 


Anomocarella falconi (King 1937) (P- 33 1 '> see also PL 9, fig 11) 

Mila Formation, Darreh Shu, Loc. 8 of King 1937 

Fig. I. Latex impression taken from a cast of the holotype, external mould of cranidium, x 4. 
Original of King 1937 • ^S- 7^> GSI 16306 (cast In 36891). 

Fig. 2. Small well-preserved cranidium retaining exoskeleton, dorsal view, x 10. In 36909a. 
Same rock fragment as cranidium, Fig. 4. 

Fig. 3. Internal mould of large cranidium, x 5. In 36908. 

Figs 4, 5, 8. Cranidium with exoskeleton, incomplete over posterior part of glabella. Dorsal 
and oblique lateral views, x 10 ; Fig. 8, detail of right side to show fine granulation, x 20. 
In 36909b. Same rock fragment as cranidium, Fig. 2. 

Koldiniella mitella Sivov 1955 (p. 333) 
Mila Formation, near Sanghabad, Alborz Mountains 

Figs 6, 7. Well-preserved cranidium in palpebral and right lateral views, x 8. It 14021. 

Figs 9, 10. Pygidium, dorsal and posterior views, x 10, showing terrace lines confined to border. 
It 13477. 

Fig. 15. Pygidium, relatively longer than previous specimen, x 10. It 14022. 

Fig. 16. Hypostome tentatively assigned to this species, ventral view, x 10. It 13506. 

Figs 13, 14. Small cranidium, possibly a small growth stage of Koldiniella mitella but may be 
referable to Pavakoldinia (see text, p. 333), x 10. It 13476. 

Tsinania ? sp. (p. 334) 
Mila Formation, near Sanghabad, Alborz Mountains 

Figs ii, 12. Incomplete cranidium, exfoliated, anterior and approximate palpebral views, 
xio. It 13493. 

Figs 17-19. Small cranidium, retaining exoskeleton, anterior and lateral views, x 7 ; Fig. 19, 
palpebral view, x 10. It 13458. 

Bull. By. Mits. nat. Hist. (Geol.) 27, 4 



Dorypyge sp. (p. 327) 

Mila Formation, near Sanghabad, Alborz Mountains 

Figs i, 2. Pygidium retaining most of exoskeleton, dorsal and oblique lateral views, x 6. 
Note impression of part of doublure posteriorly and thin subsidiary spine below third 'main' 
marginal spine on left. It 13461. 

Hadragnostus edax sp. nov. (p. 324) 
Mila Formation, near Sanghabad, Alborz Mountains 

Fig. 3. Cephalon, glabella broken, x 10. It 13468. 

Figs 4-6. Pygidium, tectonically shortened, dorsal, posterior and left lateral views, x 10. 
Note terminal node (Fig. 5). It 13472. 

Figs 7, 8, 12. Holotype pygidium, dorsal, posterior and left lateral views, x 10. Note articu- 
lating half-ring. It 13463. 

Figs 9, 10. Cephalon cut by joints, dorsal and lateral views, x 10. It 13483a, on same block 
as Figs 14, 15. 

Fig. II. Cephalon, showing rounded glabellar rear, x 10. It 14023. 

Fig. 13. Small pygidium, x 20. It 13471. 

Figs 14, 15. Cephalon, glabella damaged, dorsal and posterior views, x 15. Note faint node 
at posterior end of glabella. It 13483b, on same block as Figs 9, 10. 

Bull. Br. Mils. tiat. Hist. (Geol.) 27, 4 


Peronopsis fallax (Linnarsson 1869) aff. minor (Brogger 1878) (p. 326) 

Mila Formation, near Sanghabad, Alborz Mountains 

Figs i, 2. Large cephalon, latex cast of external mould, dorsal and lateral views, x 10. Note 
narrow axial furrow. It 13480b, on same block as Fig. 14. 

Figs 3, 4. Cephalon, slightly compressed laterally, exfoliated in region of axial furrow which 
appears wide, dorsal and lateral views, x 10. It 13480c. 

Figs 5. 8. Pygidium retaining exoskeleton, slightly deformed obliquely, dorsal and lateral 
views, X 10. It 13485. 

Fig. 6. Large pygidium, partly exfoliated, x 10. It 13467. 

Fig. 7. Pygidium, x 10. It 14025. 

Figs 9, 11. Cephalon, laterally compressed, exfoliated in region of axial furrow, lateral and 
dorsal views, x 10. Note elongate glabellar node and preglabellar depression. It 14024. 

Fig. 10. Small cephalon exfoliated in region of axial furrow, x 10. Note preglabellar de- 
pression. It 14026. 

Figs 12, 13. Pygidium, longitudinally compressed and partly exfoliated, dorsal and lateral 
views, X 10. It 13469. 

Fig. 14. Cephalon retaining exoskeleton, showing Ptychagnostus-\\\^e basal lobes, x 10. It 
i348od, on same block as Figs i, 2. 

Peronopsis fallax (Linnarsson 1869) cf. minor (Brogger 1878) (p. 327) 
Mancetter Grits and Shales, Merevale No. 3. Borehole, near Nuneaton, Warwickshire, England 

Fig. 15. Damaged internal mould, x 10. Institute of Geological Sciences, BDA 1781. 

Bull. Br. Mus. nat. Hist. (Geol.) 27, 4 





».'»^^"« ;i 


to Vol. 27 No. 4 (Miscellanea) 

New taxonomic names and the page numbers of the principal references are printed in bold 
type. An asterisk (*) denotes a figure. 

Abhar 322 

Acrospiriferinae 276 

Acuminothyris 275, 282 

Africa 309-12 

Agnostidae, Agnostinae 322-5 

Agnostoglossa simplexiformis 327 

Agnostus pisiformis Zone 323, 327 

simplexiformis 327 
Albian 301 
Alhor sella Zone 322 
Alborz Mountains 321, 324, 338 
algae 285 
A lispirifer 275-6 
Alokistocaridae 335 
America 312, 319 
Amesopleura 275 

novamexicana 278 
Androstrobus 291 
Anomocarella 322, 324, 330-1, 332 

chinensis 330, 332 

concava 332 

falconi 321, 328, 330, 331-2, 336 ; pi. 9, 
fig. II ; pi. 10, figs 1-5, 8 
Anomocarellidae 331 
Anomozamites 291 
Ardennes-London landmass 285-6 
Arizona 301 

Asaphiscidae 324, 328-34 
Auritamidae 335 
Australia 311-2,319,337; see Mindyallan 

Avon 285 

Baltagnostus 324 

Bannister, Dr K. 298 

Barbus 298 

Baringo, Lake 295-6 

Bath 285 

Bathampton Down 285-6 

Bathonian 285-6 

Battus integer 326 

Beast Cliff 291 

Belgium 282 

Bennettitaleans 291 

Bisbee Quadrangle 301 

Blastochaetetes bathonicus 285-6, 287-8 ; 

pi. 2 
Blountia 332 
BoUand 278, 281-2 


border furrows, true and false 337 
biachiopods 275-84, 286, 301-4, 321 
Brachyrhyncha 296-7 
Brachythyrina 276, 282 
Brunton, C. H. C. 275-84 
bryozoa 285-6 

California 301-4, 311 

Cambrian 321-40 

Carboniferous, Lower 275-84 

Caspian Sea 321 

Caytonia 291 

Celsifornix 275 

Chaetetida, Chaetetids 285-8 

Chal-i-Sheh 322, 336 

Chelidonocephalus 321-33, 334-6, 337-40 

alifrons 321-3, 331-2, 335, 336-7, 338 ; 
pi. 8, figs 1-5 ; pi. 9, fig. II 

preannulatus 321, 332-3, 336-7, 

338-9 ; pi. 8, figs 6-8 ; pi. 9, figs 1-5, 7 

sp. 322-3 
Chile 319 
Cirencester 285-6 
Clitheroe 278 
Cnidaria 285 
Conocoryphidae 324 
coprolites 289-94, 290* ; pi. 3, fig. 4 
coral 286 
Cotswolds 285-6 
crabs, freshwater 295-300 
Cracow 303 
Crepicephalus Zone 323 
Cretaceous 301-4 
Croiset collection 308 
Cyclothyridinae 302-3 
Cyclothyyis 301-4 

americana 301 

antidichotoma 303 ; pi. 5, figs 2-3 

densleonis 301, 302*, 302-3 ; pi. 5, figs i, 


latissima 303 

mirabilis 303 

subtrigonalis 301 
Cyrtia 276 
Czekanowskia 291 

Darreh Shu 322, 331 
Davidson collection Z'j'j-S 
Davies, Dr R. G. 321-2 



Deltaic, Lower 289 

Delthyrididina 276 

Devonian, Upper 282 

Dictyella 324 

Dictyites 324 

Dorypyge 322-3, 327, 333, 338 ; Zone 322-3 

aenigma 333 

iranensis 328 

khademi khademi 328 
papulosa 328 

richthofeni 327 
laevis 328 
laiwuensis 328 
n. subsp. aff. richthofeni 327 

sp. 321, 323, 327-8 ; pi. 11, figs 1-2 
Dorypygidae 327-8 
Dovedale 277, 279-80 
Drepanura Zone 322-3 

{Spinopanura) 322 
Dublin 282 

echinoderms 286 

Eleutherokomma 275-6 

Elliott, G. F. 285-8 

England 301, 303 

eretes Zone 323 

Europe 301 

Eymekops rectangularis 335 

Faringdon 303 
Fermanagh 278-81 
fish 291, 298 
flamingos 305-14 
Florida 311 
Forest Marble 286 
FoRTEY, R. A. 321-40 
France 285, 303, 305, 307, 309 
Fusella 275, 276-8, 279-84 

convoluta 281-2 

fusiformis 275-7, 278-82; pi. i, figs 5- 
12, 20-21 

llanoensis 278 

rhomboidea 275, 277-8, 280*, 281*, 281- 
2 ; pi. I, figs 13-19 

roemerianus 282 

tornacensis 277 

triangularis 282 

trigonalis 281-2 

Germany 309 
Gervaisia 307-8, 308-9 

croiseti 305, 306*, 307*, 308, 310 
Gilbertson collection 278, 282 
Gloucestershire 285 

Grandioculus falconi 330 

megalurus 330 
Gristhorpe Bed 291 

Haanakdar 322 
Hadragnostus 324 

edax 321, 323, 324-5 ; pi. 11, figs 3-15 

las 323-5 

modestus 323-5 
Harrison, C. J. O. 305-20 
Hartshillia 324 
herbivores 289 
Hill, C. R. 289-94 
Hinde collection 285 
Holloway, Miss M. L. 312, 319 
Horsetown Group 301 
Hundwarella 328 
Hydrozoa 285 
Hypermecaspis 328 

Illaenidae 333 
Illaenuridae 333 
Illaenurus 333 

caneus 334 
Iniotoma 328 
Inouyella 335 

peiensis 335 
insects 291 
Iran 321-40, 322* 
Irania 328 

falconi 328, 330-1 

pisiformis 328-9 
Iranoleesia 322, 324, 328-9, 330 ; Zone 

falconi 321, 328, 330 

pisiformis 321-2, 328, 329-30; pi. 9, 
figs 6, 8-10, 12 

Jell, Dr P. 
Jurassic 285-94 

Kabarsero 295 

Kalimale 295-6 

Kamwina 295-6, 298 

Kaolishania Zone 322 

Kashmir 328 

Keates, C. 319 

Kenya 295-300, 305, 309, 311 

Kingstonia 333 

Koldinia 324 

Koldiniella 322-3, 332-3, 338 

convex a 333 

mitella 321-3, 329, 332, 333-4; pi. 10, 
figs 6-7, 9-10, 13-16 

prolix a 333 



Kootenia 328 
Kulyumbe 323 

laevigata Zone 323 
Leakey, Dr L. S. B. 311, 317 
Leighton Buzzard 303 
Leistegiidae 335 
Lejopyge laevigata Zone 323 
Licharewiinae 276 
Limekiln Pit 285 
Liopeishania 333 

lubrica 333 

spannensis 333 
Lirrangopotamonautes see Potamonautes 
Llanoaspis walcotti 335 

Macelloura 333 

Ma'dan 322, 329 

mammal 291-2 

Mapan Formation 332 

Mapania 331 

Mapaniidae 335 

Maryvillia 332 

Meneviella 324 

Mexico 301, 312 

Mila Formation 321, 323-4, 331, 338 

zones 322 
Mila Kuh 322 

Mindyallan Stage 323, 328, 337 
Miocene 295-300, 305-14, 319 
Mirifusella 277 
molluscs 286 
Montana 323 

Morris, S. F. 295-300, 321 
Mucrospiriferidae 275-82 
Mural Limestone 301 

Nails worth 285 
Nairobi 296 
Namanoia 335 
Namanoiidae 335 
New Mexico 278 
New South Wales 278 
Ngorora Formation 295-300 
Nilssoniopteris vittata 291 
Nuneaton 323 

Olduvai 315-20 
Oligocene 305-14 
oncolites 285 
Ordosia 335 

fimhricauda 335 
ornithopod dinosaur 291 
Orthopotamonautes see Potamonautes 

ostracods 295 
Owen, E. F. 301-4 

Paeckelmanella 2.'jb, 282 
Paeckelmanellidae 276 
Papyriaspididae 335 
Parakoldinia 333 ; pi. 10, figs 13-14 

plana 334 

? sp. 321 
Paris Basin 285 

Pedinocephalina - Toxotis (?) Zone 323 
Peichishania 335 

rectangularis 335 
Peishania 333 

lubrica 333 
Pelecani 315 
Pelecanidae 315 
Pelecaniformes 315 
Pelecanus 315-19 

aethiopicus 315-19, 316* ; pi. 7 

conspicillatus 318-19 

crispus 316*, 318-19 

erythrorhynchus 318-19 

gracilis 319 

grandiceps 319 

halieus 319 

intermedius 319 

occidentalis 318-19 
thagus 319 

odessanus 319 

onocrotalus 316*, 318-19 

proavus 319 

rufescens 316*, 318 
pelican 315-20 
Perignat 308 
Permian 282 
Peronopsis 324, 326 

fallax ferox 326 

aff. minor 321, 323, 326-7 ; pi. 12, 

figs I -14 
cf. minor 323, 326 ; pi. 12, fig. 15 

integer 326 
Peru 319 
Phoeniconaias 305, 308-11 

floridanus 311 -12 

gracilis 311 

minor 305, 306*, 307*, 309, 309*, 311 

novaehollandiae 311 

stocki 312 
Phoeniconotius eyrensis 312 
Phoenicoparrus 305, 308, 310 

andinus 305 

jamesi 305, 306*, 307*, 309* 
Phoenicopteridae 310 



Phoenicopterus 305, 308, 310 

aethiopicus 305, 309*, 310*, 311-12 
pi. 6 

copei 312 

croiseti 305, 307-8 

minuius 311 

ruber 305, 307*, 308-11 
antiquorum 305 

chilensis 305, 306*, 307*, 309*, 310* 
ruber 305 
Pickford, Dr M. 295 
plant cuticles 289-92 
Pleistocene 311-2, 315-20 
Plethometopus convergens 334 

obtusus 334 
Plethopeltoides lepidus 334 
Pliocene 31 1-2, 319 
Poshania 335 

poshanensis 335 
Poshauia 335 
Potamidae 296-7 

Potamon {Potamonautes) lirrangensis 296 
Potamonautes 296, 297-8 

lirrangensis 296 

per lata 296 

[Lirrangopotamonautes) 295, 296, 297 
lirrangensis 296, 298 
tugenensis 295, 296-7, 298 ; pi. 4 

{Orthopotamonautes) 295, 297 
depressus 298 
johnstoni 298 
Prochuangia Zone 322 
Protohedinia 329 
Pterophyllum 291 
Pterospirifer -zj^-b, 282 
Ptilophyllum 289-94 

pectinoides 290*, 291 ; pi. 3 
Ptychagnostus 327 

atavus 326 
ptychoparioids 321, 323 
Puy-de-D6me 307 
pygostyle 297 

Qanli-Chapoghlu 322 
Qazvin 321 

Quadragnostidae 326-7 
quasivespa Zone 323 
Queensland 323 

rabbits 289, 291 
reptiles 291 
Rhynchonella 301 

densleonis 301-2 
Rhynchonellidae 301-3 

Rhyssometopacea 335-7 

Rhyssometopus princeps 337 

Rift Valley 298 

RissoNE, A. 275-84 

Roseberry Topping plant bed 289-92 

RuSHTON, A. W. A. 321-40 

Rusinga Island 305, 309, 311 

Salair 323 

Sanghabad 321-2, 324, 338 

Saukia Zone 322 

Scandinavia 323 

Schulze- Ammonia maceration 291 

Sclerospongiae 286 

Shahmirzad 322 

Shasta County 301 

sheep 291-2 

Shiraz 321 

Siberia 323 

S olenites 291 

Spinopanura see Drepanura 

Spirifer 276 

bicarinata 276, 282 

convoluta 282 

cuspidatus 281-2 

distans 282 ; pi. i, figs 1-4 

fusiformis 276, 278 

mundula 278 

novarnexicana 278 

rhomboidea 277 

rhomboides 276 

roemerianus 282 

tornacensis 277 

triangularis 282 
Spiriferacea 275-82 
Spiriferidae 275-6 
Spiriferidina 276 
Spirifinaella 276 
sponges 286 
stromatoporoids 285 
Strophopleura 275, 282 

anterosa 278 
Strophopleurinae 275-6, 277-82 
Syringothyridacea 276 
Syringothyrididae 277 
Syringothyris 281-2 

cuspidatus 281-2 

tabulates 285 
Taitzuia 335 
insueta 335 
Taleghan Range 324, 338 
Tang-i-Tehbud Bezuft 322 
Tanout 298 



Tanzania 315, 317 
Tehran 321 
Tengfengiidae 329 
Tennessee 335 
Terebratulidae 301 
Thelphusa perlata 296 
Thomas, Dr H. H. 289 
Tiltup's End 285-6 
Tomes collection 285 
Tournaisian 282 
Toxotis see Pedinocephalina 
Treak Cliff 281-2 
trilobites 321-40 

effaced 324, 332-3 
Tsinania 324, 334 

caneus 334 

? sp. 321, 323, 334; pi. 10, figs 11-12, 
Tsinaniidae 324, 333 
Tugen Hills 295-6 

Unispirifer 277-8 

Victoria, Lake 309, 311 

Visean 279-82 

Voiseyella 275-6, 278, 282 

anterosa 278 

novamexicana 278 

Walker, C. A. 305-20 
Wanwanoglohus convexus 334 
Williamsonia 292 

hildae 291 

leckenhyi 291 
Witchell collection 285 

Xanthoidea 296-7 

Yorkshire 289-94 

Zagros Thrust 321 





1. Cox, L. R. Jurassic Bivalvia and Gastropoda from Tanganyika and Kenya. 
Pp. 213 ; 30 Plates ; 2 Text-figures. 1965. out of print. 

2. El-Naggar, Z. R. Stratigraphy and Planktonic Foraminifera of the Upper 
Cretaceous — Lower Tertiary Succession in the Esna-Idfu Region, Nile Valley, 
Egypt, U.A.R. Pp. 291 ; 23 Plates ; 18 Text-figures. 1966. £11. 

3. Davey, R. J., DowNiE, C, Sarjeant, W. a. S. & Williams, G. L. Studies on 
Mesozoic and Cainozoic Dinoflagellate Cysts. Pp. 248 ; 28 Plates ; 64 Text- 
figures. 1966. £8.20. 

3. Appendix. Davey, R. J., Downie, C, Sarjeant, W. A. S. & Williams, G. L. 
Appendix to Studies on Mesozoic and Cainozoic Dinoflagellate Cysts. Pp. 24. 
1969. 95p. 

4. Elliott, G. F. Permian to Palaeocene Calcareous Algae (Dasycladaceae) of 
the Middle East. Pp. iii ; 24 Plates ; 16 Text-figures. 1968. out of print. 

5. Rhodes, F. H. T., Austin, R. L. & Druce, E. C. British Avonian (Carboni- 
ferous) Conodont faunas, and their value in local and continental correlation. 
Pp- 313 J 31 Plates ; 92 Text-figures. 1969. ;^ 

6. Childs, a. Upper Jurassic Rhynchonellid Brachiopods from Northwestern 
Europe. Pp. 119 ; 12 Plates ; 40 Text-figures. 1969. £5.25. 

7. Goody, P. C. The relationships of certain Upper Cretaceous Teleosts with 
special reference to the Myctophoids. Pp. 255 ; 102 Text-figures. 1969. 


8. Owen, H. G. Middle Albian Stratigraphy in the Anglo-Paris Basin. Pp. 
164 ; 3 Plates ; 52 Text-figures. 1971. £7.20. 

9. SiDDiQUi, Q. A. Early Tertiary Ostracoda of the family Trachyleberididae 
from West Pakistan. Pp. 98 ; 42 Plates ; 7 Text-figures. 1971. £9-60. 

10. Forey, p. L. a revision of the elopiform fishes, fossil and Recent. Pp. 222 ; 
92 Text-figures. 1973. ;^ii.35- 

11. Williams, A. Ordovician Brachiopoda from the Shelve District, Shropshire. 
Pp. 163 ; 28 Plates ; 11 Text-figures ; no Tables. 1974. £12.80. 

Printed in Great Britain hy John Wright and Sons Ltd. at The Stonebridge Press, Bristol BS4 5NU 








GEOLOGY Vol. 27 No. 5 

LONDON: 1977 

: J JAK i977 





University of Oxford 


South African Museum, Cape Town 

Pp. 347-380 ; 15 Plates ; 9 Text-figures 

GEOLOGY Vol. 27, No. 5 

LONDON: 1977 


(natural history), instituted in 1949, is 
issued in five series corresponding to the Scientific 
Departments of the Museum, and an Historical series. 

Parts will appear at irregular intervals as they 
become ready. Volumes will contain about three or 
four hundred pages, and will not necessarily be 
completed within one calendar year. 

In 1965 a separate supplementary series of longer 
papers was instituted, numbered serially for each 

This paper is Vol. 27, No. 5 of the Geological 
{Palaeontological) series. The abbreviated titles of 
periodicals cited follow those of the World List of 
Scientific Periodicals. 

World List abbreviation : 
Bull. Br. Mus. nat. Hist. (Geol. 

ISSN 0007-1471 

Trustees of the British Museum (Natural History), 1977 


Issued 2y January igyy Price £6-10 





Abstract .... 
I. Introduction 

Location of Specimens . 
Field Localities . 
Dimensions of Specimens 
II. Systematic palaeontology . 
Genus Phylloceras 
Subgenus Phylloceras 

Phylloceras {Phylloceras) serum 
Subgenus Hypophylloceras . 

Group of Phylloceras (Hypophylloceras) thetys 

Phylloceras [Hypophylloceras) subalpinum ellipticxmi 
Phylloceras {Hypophylloceras) aphrodite . 
Group of Phylloceras (HypopJiylloceras) velledae . 
Phylloceras {Hypophylloceras) velledae . 

Phylloceras {Hypophylloceras) velledae velledae 
Phylloceras {Hypophylloceras) velledae infiatum 
Phylloceras {Hypophylloceras) sp. aff. velledae . 
Group of Phylloceras {Hypophylloceras) seresitense 
Phylloceras {Hypophylloceras) seresitense 

Phylloceras {Hypophylloceras) seresitense seresitense 
Phylloceras {Hypophylloceras) androiavense 
Phylloceras {Hypophylloceras) woodsi woodsi 
Phylloceras {Hypophylloceras) mikobokense 
Genus Partschiceras .... 
Partschiceras umzambiense . 
Parschiceras sp. juv. ? cf. baborense 
Genus Carinophylloceras 

Carinophylloceras collignoni . 

III. Acknowledgements .... 

IV. References ...... 

V. Index ....... 

VI. Erratum ...... 



Phylloceratid ammonites are present in rocks of Barremian, Albian, Cenomanian, Santonian, 
Campanian and Maastrichtian ages in Zululand and Natal, South Africa. They form a sub- 
stantial proportion of the normally coiled ammonite fauna during the Barremian and Albian, 
but are rare in Aptian and post-Albian faunas. The following forms are described below. 


Phylloceras [Phylloceras) serum (Oppel), P. {Hypophylloceras) subalpinum (d'Orbigny) 
ellipticum Kossmat, P. {Hypophylloceras) aphrodite Fallot & Termier, P. {Hypophylloceras) 
velledae velledae (Michelin), P. {Hypophylloceras) velledae (Michelin) inflatum Collignon, 
P. {Hypophylloceras) sp. aff. velledae (Michelin), P. {Hypophylloceras) seresitense seresitense 
Pervinquiere, P. {Hypophylloceras) androiavense Besairie, P. {Hypophylloceras) woodsi woodsi 
van Hoepen, P. {Hypophylloceras) mikobokense Collignon, Partschiceras umzambiense (van 
Hoepen), Partschiceras sp. juv. cf. baborense (Coquand) and Carinophylloceras collignoni Klinger, 
Wiedmann & Kennedy. 

Eight of these species are new records from the area. 


The Phylloceratidae are the root stock of the Jurassic and Cretaceous phyllo- 
ceratids, appearing in the Lower Jurassic (Sinemurian) and ranging to the Upper 
Cretaceous (Lower Maastrichtian) . They have a world-wide distribution, but for 
most of the Cretaceous they are typically rare in the boreal region of north-western 
Europe and in the western interior of North America, In South Africa, species 
are known from the Barremian to Lower Maastrichtian, and are relatively common 
in the Barremian and Albian. 

The following species are described below : 

Phylloceras {Phylloceras) serum (Oppel) 

P. {Hypophylloceras) subalpinum (d'Orbigny) ellipticum Kossmat 

P. {Hypophylloceras) aphrodite Fallot & Termier 

P. {Hypophylloceras) velledae velledae (Michelin) 

P. [Hypophylloceras) velledae (Michelin) inflatum Collignon 

P. {Hypophylloceras) sp. aff. velledae (Michelin) 

P. [Hypophylloceras) seresitense seresitense Pervinquiere 

P. [Hypophylloceras) androiavense Besairie 

P. [Hypophylloceras) woodsi woodsi van Hoepen 

P. [Hypophylloceras) mikobokense Collignon 

Partschiceras umzambiense (van Hoepen) 

Partschiceras sp. juv. cf. baborense (Coquand) 

Carinophylloceras collignoni Klinger, Wiedmann & Kennedy 

Location of Specimens 

The following abbreviations are used to indicate the repositories of the material 
studied : 

BM(NH) British Museum (Natural History), London 

EMP Ecole des Mines, Paris 

MHNP Museum d'Histoire Naturelle, Paris 

SAM South African Museum, Cape Town 

SAS South African Geological Survey, Pretoria 

TM Transvaal Museum, Pretoria 

UPE University of Pretoria, Pretoria 


Field Localities 

Outline details of field localities referred to in this paper are given by Kennedy & 
Klinger (1975) ; full descriptions of sections are deposited in the Palaeontology 
Library of the British Museum (Natural History). 

Dimensions of Specimens 
All dimensions given below are in millimetres : 

D = Diameter 
Wb = Whorl breadth 
Wh = Whorl height 

U = Umbilicus 

Figures in parentheses are dimensions as a percentage of the total diameter. 


Class CEPHALOPODA Cuvier 1797 

Subclass AMMONOIDEA Zittel 1884 

Order PHYLLOCERATIDA Arkell 1950 

Superfamily PHYLLOCERATAGEAE Zittel 1884 

Family PHYLLOCERATIDAE Zittel 1884 

Subfamily PHYLLOCERATINAE Zittel 1884 

Genus PHYLLOCERAS Suess 1865 

Type species. Ammonites heterophyllus J. Sowerby 1820. 

Discussion. An extensive and useful review of the scope and interpretation of 
Phylloceras and its subgenera has been given by Wiedmann in a series of publications 
(1962a, b, 1964; Wiedmann & Dieni 1968). He stresses the slight differences of 
both species and genera within the phylloceratids, and regards many of the taxa 
proposed by previous workers as of little value on either phylogenetic or strati- 
graphic grounds. With minor reservations, his conclusions are accepted here, and 
they clarify the relations between the bewildering host of taxa referred to the group. 
Our following discussions thus rest heavily on his work, but are included here as a 
review of work otherwise available only in German. 

The genus Phylloceras is thus typified by the group of Phylloceras heterophyllum. 
Rhacoceras Hyatt 1867 and Heterophylloceras Kovacs 1939 are both objective 
synonyms, while the poorly defined Xeniophylloceras Buckman 1921, Hantkeniceras 
Kovacs 1939 and Pseudophylloceras Beznosov 1958 do not bear separation on either 
gross morphological features or sutural details. Furthermore, fasciculate ribbing 
occurs in some Phylloceras s.s., suggesting that the Pseudophylloceratinae (Beznosov 
1958) - genera such as Neophylloceras Shimizu 1934 and Epiphylloceras Collignon 
1956 - do not bear separation from Phylloceras s.s. The following subgenera can 
be recognized. 



Phylloceras Suess 1865, type species Ammonites heterophyllus J. Sowerby 1820, 
Lower Jurassic to Lower Cretaceous. 

Zetoceras Kovacs 1939, type species Ammonites zetes d'Orbigny 1850, Lower to 
Middle and ? Upper Jurassic. 

Geyeroceras Hyatt 1900, type species Ammonites cylindricus J. de C. Sowerby 
1 83 1, Lower Jurassic. 

Hypophylloceras Salfeld 1924, type species Phylloceras onoense Stanton 1895, 

Subgenus PHYLLOCERAS Suess 1865 

Phylloceras (Phylloceras) serum (Oppel) 

(Fig. I ; Pis 1-2 ; PI. 3, figs 1-2) 

1865 Ammonites serus Oppel : 550. 

1868 Phylloceras serum Oppel ; Zittel : 66, pi. 7, figs 5, 6. 

1885 Phylloceras semistriatum d'Orbigny ; Neumayr : 83, pi. i, fig. 2 (non d'Orbigny). 

1890 Phylloceras serum Oppel ; Toucas : 574, pi. 13, fig. 2. 
? 1891 Phylloceras cf. thetys d'Orbigny ; Sayn : 141, pi. i, fig. i. 

1901 Phylloceras serum Oppel var. perlobata Sayn : 7, text-fig. 3, pi. i, figs 6-8. 

1901 Phylloceras thetys d'Orbigny ; Sayn : 36, pi. i, fig. 3 only. 

1907 Phylloceras serum Oppel ; Pervinquiere : 12, pi. i, fig. 3. 

1914 Phylloceras serum Oppel var. perlobata Sayn ; Zwierzycki : 33, pi. 4, figs 2, 3. 

1949a Phylloceras spathi Collignon : 63, pi. 9, figs i-ib. 
? 1949a Phylloceras rohionense Collignon : 64, 65, text-figs i-ia. 

1 95 1 Phylloceras serum (Oppel) ; Arnould-Saget : 5, pi. i, fig. 5. 

1957 Hyporbulites betieri Busnardo ; Busnardo & David : 84, pi. i, figs 1-2 ; text-figs 4, 5. 

i960 Euphylloceras serum Oppel ; Drushchitza & Kudryavtseva : 251, pi. i, fig. 8. 

i960 Phylloceras serum Oppel ; Collignon : pi. 134, figs 506, ? 507. 

1962a Phylloceras serum Oppel ; Collignon : 20, pi. 181, fig. 817 ; 33, pi. 186, fig. 853 ; 76, 
pi. 214, fig. 916. 
? 1962a Phylloceras spathi Collignon ; Collignon : 74, pi. 203, fig. 915 ; pi. 204, fig. 915a. 

1966 Phylloceras cf. perlobatum Sayn ; Rawson : 455, pi. 72, figs 1-3. 
? 1968 Ph. {Phylloceras) spathi Collignon ; Wiedmann & Dieni : 22. 

1968 Ph. {Phylloceras) serum (Oppel) ; Wiedmann & Dieni : 20. 

Lectotype. The specimen figured by von Zittel (1868) as his pl. 7, fig. 5 is 
herein designated lectotype. 

Material. Fourteen specimens, all from the Upper Barremian (Barremian I), 
Loc. 170, Mlambongwenya Spruit, northern Zululand. BM(NH) C78652-61. 
SAS LJE 132b, 54/19, 54/33, Z19. 







Zittel 1868 : 66 





Collignon i960 : 

pl. 134, fig. 566 


35 (36) 

57 (59) 


Collignon 1962a : 

pl. 204, fig. 916 


43-0 (35) 

72-0 (59) 


BM(NH) C78657 


28-2 (33) 

49-4 (57) 


6-4 (7) 

BM(NH) C78660 


44-5 (37) 

70-8 (59) 


8-6 (7) 



Fig. I. 

External suture line of Phylloceras (Phylloceras) serum Oppel. 


SAS LJE 132b. 

Phylloceras spathi, from CoUignon 1949a : 62. 

Example i ii2-o 46-0 (41) 63-0 (56) 073 6-o (5) 

Example 2 iio-o 44-0 (40) 65-0 (59) 0-67 6-o (5) 

Example 3 113-0 40-0 (35) 66-o (53) o-6o 6-o (5) 

Phylloceras rohionense, from CoUignon 1949a : 64. 

Holotype 141-0 45-0 (32) 8i-o (57) 0-55 5-0 (4) 

Description. The coiling is very involute with a tiny, deep umbilicus measuring 
7% of the diameter. The umbilicus is undercut, the umbilical wall rounded, and 
the umbilical shoulder gently sloped, so that there is a shallow conical depression 
surrounding a deep umbilical pit. The whorl section is a compressed oval, with 
flattened subparallel lower flanks and upper flanks which converge towards broadly 
rounded shoulders and a broad, rounded venter. 

The shell surface is ornamented by fine, dense, flexuous striae. These arise at the 
umbilical seam and are very fine, dense and crowded over the umbilical wall, 
shoulder and lower flank, strengthening on the outer flank and venter. They pass 
straight across the lower flank, flex slightly backwards some way below mid-flank, 
to sweep gently forwards over the upper flank and shoulder. They pass across the 
broad venter with a faint forwards projection. Internal moulds bear a fainter but 
comparable ornament. 

The suture line is complex, phylloid, the saddles having tetraphylloid terminations, 
as shown in Fig. i. 

Discussion. Flattened sides, broad venter, flexuous ornament and tetraphylloid 
saddles clearly place our specimens within the range of variation ascribed to 
Phylloceras {Phylloceras) serum by CoUignon (1962), Arnould-Saget (1951) and 
Wiedmann & Dieni (1968), whom we foUow in regarding Sayn's variety perlohata 



and the ' Hyporbulites' hetieri of Busnardo as synonyms. Phylloceras {Phylloceras) 
spathi Collignon is very close to P. (P.) serum in ornament, and overlaps in relative 
proportions, the large specimen figured by Collignon (1962a : pi. 203, fig. 915) having 
a similar suture, which suggests that this species may also fall into synonymy. 
Wiedmann & Dieni (1968 : 22), however, state that P. (P.) spathi lacks ornament on 
internal moulds and differs from P. (P.) serum mainly in whorl shape. Phylloceras 
rohionense, from the Barremian of southern France (Basses-Alpes) , may also be a 

P. (P.) serum has, on occasion, been confused with Phylloceras of the thetys group, 
with which it is homoeomorphic ; these forms have, however, diphylloid saddle 
terminations as do forms such as Phylloceras sablyense Karakash. 

Occurrence. Phylloceras {Phylloceras) serum has a long time range and a wide 
geographic distribution, which may be summarized as follows : 

Upper Jurassic (Tithonian) of central Europe (Stramberg Limestone), southern 
France, north Africa (Tunisia) and Madagascar ; Lower Cretaceous of southern 
France (Valanginian and ? Barremian), Sardinia ( Valanginian) , Madagascar 
(Hauterivian and Valanginian) and Zululand (Barremian). 

Subgenus HYPOPHYLLOCERAS Salfeld 1924 

Type species. Phylloceras onoense Stanton 1895. 

Diagnosis. Compressed, discoidal involute phylloceratids, inner whorls occasion- 
ally constricted {'Aphroditiceras', ' Euphylloceras' , ' Goretophylloceras') . Ornament 
of both shell and moulds consists of weak radial striae, strengthening on the venter ; 
occasionally fasciculate in the umbilical region {'Neophylloceras' , ' Epiphylloceras') . 
Suture with diphylloid {'Goretophylloceras'), tetraphylloid {'Hyporbulites') or poly- 
phylloid saddles (from Wiedmann 1964 : 173). 

Discussion. Following Wiedmann, we regard Neophylloceras Shimizu 1934, 
Paraphylloceras Shimizu 1935, Hyporbulites Breistroffer 1947, Goretophylloceras 
Collignon 1949a, Euphylloceras Drushchitz 1953 and Aphroditiceras Mahmoud 1956 
as synonyms of Hypophylloceras. The group thus represents a variable cluster of 
phylloceratids linked by their apparent common origin in Phylloceras of the serum 
group during the early Cretaceous ; detailed study shows that there is every inter- 
mediate between the supposedly diagnostic morphologies of the various genera 
placed in synonymy. Furthermore, the intimate gradational relationship between 
P. (P.) serum and some Hypophylloceras species suggests that even subgeneric 
separation is more of a stratigraphic convenience than of phylogenetic value. 

Occurrence. Hypophylloceras has a world-wide distribution, although rare in 
the boreal region and the western interior of North America. Species first appear 
in the Valanginian, and range to the Lower Maastrichtian. 

I. Group of P. {Hypophylloceras) thetys (d'Orbigny) 

This species group is characterized by di- or triphylloid saddles throughout 
ontogeny. It is thus separable from the velledae group where saddles are at first 


diphylloid, becoming tetraphylloid when adult, and the seresitense group where 
saddles are tetraphylloid throughout ontogeny. The following are referred to the 
group : 

1. Hypophylloceras thetys (d'Orbigny) (1841 : 174 ; pi. 53, figs 7-9), Valanginian 
to Barremian. 

2. Hypophylloceras thetys (d'Orbigny) diegoi Boule, Lemoine & Thevenin (1906 : 10 ; 
pi. I, figs 5, 7 ; text-fig. 4), Barremian to Cenomanian. 

3. Hypophylloceras thetys (d'Orbigny) majoricense Wiedmann (1964 : 178 ; pi. 15, 
fig. I ; pi. 18, fig. 3 ; text-fig. 37), Aptian to Albian. 

4. Hypophylloceras sablyense Karakash (1907 : 39 ; pi. 3, figs I2a-b ; pi. 14, 
figs 8a-b ; pi. 24, fig. 4), Barremian. 

5. Hypophylloceras fortunei (Honnorat-Bastide 1892) (Wiedmann 1964 : 192), 

6. Hypophylloceras aptiense aptiense Sayn (1920 : 195 ; pi. i, fig. 4 ; text-fig. 2), 

7. Hypophylloceras aptiense Sayn applanatum Wiedmann (1964 : 190 ; pi. 15, 
fig. 5 ; text-fig. 43), Aptian. 

8. Hypophylloceras moreti Mahmoud (1955 : 76 ; pi. 5, figs 2-4 ; text-fig, 44), 

9. Hypophylloceras aphrodite Fallot & Termier (1923 : 25 ; pi. 2, figs 5-7 ; text-fig, 
5), Albian. 

10. Hypophylloceras subalpinum subalpinum d'Orbigny (1841 : 283 ; pi. 8^, figs 
1-3), Albian. 

11. Hypophylloceras subalpinum d'Orbigny ellipticum Kossmat (1895 : 11 ; pi. i, 
fig. 2 ; pi. 6, fig. i), ? Aptian, Albian to Cenomanian. 

12. Hypophylloceras improvisum (Stoliczka) (1865 : 113 ; pi. 58, figs 4-4b), Ceno- 

13. Hypophylloceras subseresitense Wiedmann (1964 : 202 ; pi. 14, fig. 9 ; pi. 15, 
fig. 6 ; text-fig. 47), Aptian. 

14. Hypophylloceras sardoum Wiedmann & Dieni (1968 : 24 ; pi. i, figs i, 2 ; pi. 2, 
fig. 2 ; text-fig. 3), Albian. 

Phylloceras (Hypophylloceras) subalpinum (d'Orbigny) ellipticum Kossmat 
(Figs 2-3 ; PL 4, figs 1-4 ; PI. 5, figs 2-3 ; PI. 12, fig. 2) 

1865 Ammonites suhalpinus d'Orbigny ; Stoliczka : 114 ; pi. 58, figs 3a-c (non d'Orbigny). 
? 1877 Ammonites velledae Michelin ; Simonovich, Batsevich & Sorokin : pi. 3, fig. 3 (non 
Michelin) . 

1895 Phylloceras ellipticum Kossmat : 107 ; pi. 15, figs 2a-b ; pi. 20, figs la-b. 
non 1897 Phylloceras ellipticum Kossmat ; Parona & Bonarelli : 78 ; pi. 10, fig. 7 [= Phyllo- 
ceras aphrodite Fallot & Termier]. 

1907a Phylloceras cf. ellipticum Kossmat ; Crick : 169 ; pi. 10, figs 12 -12a. 

1907 Phylloceras ellipticum ? Kossmat ; Pervinquiere : 51 ; pi. 3, figs i, 2a-b ; text- 
fig. 4. 

1910 Phylloceras ellipticum Kossmat ; Pervinquiere : 10. 

1928 Phylloceras cf. ellipticum Kossmat ; CoUignon : 147 ; pi. i, figs lo-ioa. 


Fig. 2. External suture line of Phylloceras {Hypophylloceras) subalpinum ellipticum 
Kossmat. SASA1116, x 6. 

1943 Phylloceras japonicum Matsumoto : 674 (nom. nud.). 
1953 Phylloceras japonicum Matsumoto : 147 ; text-fig. 10 (nom. nud.). 
non 1956 Phylloceras cf. ellipticum Kossmat ; Collignon : 15 ; pi. i, figs i, la, ib 

[= Phylloceras improvisum (Stoliczka)]. 
1959 Partschiceras ? japonicum (Matsumoto) ; Matsumoto : 52 ; pi. 12, figs la-c, 2a-b, 

3a-c ; pi. 28, figs 2a-d ; pi. 29, figs la-b ; text-fig. 2. 
1964 Ph. (H.) subalpinum ellipticum Kossmat ; Wiedmann : 197 ; pi. 13, figs 5a -b ; 

pi. 14, fig. 8 ; pi. 15, figs 3a-c. 
1968 Ph. {Hypophylloceras) subalpinum ellipticum Kossmat ; Wiedmann & Dieni : 23 ; 

pi. I, fig. 4 ; pi. 2, figs la-b. 

HoLOTYPE. Kossmat's original specimen from the Utatur Group of southern 
India (1895 : pi. 15, figs 2a-b ; pi. 20, fig. i). 

Material. We have fourteen specimens : BM(NH) C18139 (figured by Crick 
1907a) probably from the Cenomanian of the Skoenberg region, BM(NH) C78662 
from Bed 6, Loc. 51, BM(NH) C78664 from Loc. 54, SAS 2477 and Z38 from Loc. 57, 
all from the Mzinene River (Albian V) ; BM(NH) C78663 from Bed 4, Loc. 64, 
BM(NH) C78665, SAS 945, 66/Pi from Loc. 66, Munywana Creek (Albian V), 
BM(NH) C78666, SAS A499, 10822 and A945 from Loc. 62, the Skoenberg, Mzinene 
River (Cenomanian II-III) and SAS An 16 from the Albian of the Mzinene (precise 
horizon unknown). BM(NH) C78686 from Bed 4, Loc. 64, on the Mun3rwana Creek, 
may also belong here. 







Holotype (from 

Kossmat 1895) 


28-5 (40) 

41-0 (58) 


5-0 (7) 

BM(NH) 78662 


17-8 (40) 

25-3 (56) 


3-3 (7) 

BM(NH) C78669 


20-0 (34) 

35-2 (60) 


3-6 (6) 

SAS 10822 


17 (30) 

31 (55) 


5-0 (9) 

SAS Z447 


20 (34) 

35 (59) 


4-0 (7) 

SAS 66/Pi 


17-5 (31) 

31 (55) 


4-5 (8) 


Description. The coiling is very involute, with a tiny, funnel-shaped umbiHcus 
(6-9% of diameter). The whorl section is a compressed oval, the whorl breadth 
to height ratio varying from 0-55 to 0-71, the greatest breadth being typically at 
or just below mid-flank. The umbilical shoulder slopes gently outwards, producing 
a funnel-shaped umbilical depression, the flanks being gently inflated, converging 
to a narrow rounded venter. The surface of the shell is ornamented by low, flat, 
radial folds which arise at the umbilical seam as mere striae, but strengthen across 
the lower flank, flexing faintly forwards as they do so, to flex faintly back at mid- 
flank and pass almost straight across the upper flank and venter. The internal 
mould is smooth, or bears only faint, low folds. 

The suture is moderately subdivided with triphylloid saddles (Figs 2-3). 

Discussion. Ornament and whorl section clearly place our specimens close to 
P. {Hypophylloceras) subalpinum ellipticum ; specimens figured as PI. 5, figs 3a-c 
and PI. 6, figs 2a-b have the identical weak fold-like ribs and elliptical whorl section 
clearly shown in Kossmat's figures and in material illustrated recently by Wiedmann 
(1964). Other specimens, more compressed than previously-described material 
referred to this species (whorl breadth to height ratio as low as 0-55) but otherwise 
identical, we also place within the range of subalpinum ellipticum. None of our 
specimens show the constrictions described by Kossmat, nor the fine striations 
developed on well-preserved material (e.g. Wiedmann 1964 : pi. 15, fig. 3c). The 
dimensions of ellipticum overlap those of P. {H.) subalpinum subalpinum, and 
involution and sculpture are identical. These similarities led Stoliczka to refer 
his Indian material to P. (H.) subalpinum s.s. Kossmat, however, separated the 
Indian material on the basis of the elliptical whorl section with the greatest breadth 
at or about mid-flank, and the presence of constrictions and fine striations in 
ellipticum, as opposed to the trigonal whorl section in subalpinum. As Wiedmann 
(1964 : 198) has noted, no body chambers of subalpinum have been described, so 
that the constrictions and fine striations cannot be used to separate these forms. 
Whorl section remains as one distinguishing character, while the suture lines of the 

Fig. 3. External suture line of Phylloceras {Hypophylloceras) subalpinum ellipticum 

Kossmat. SAS Z447, x 3. 


present material show that virtually all the saddles are subtriphylloid, thus 
substantiating Wiedmann's (1964) suggestion that the greatest difference between 
the two subspecies is in the configuration of the saddles, especially L/Ug. 

' Partschiceras' japonicum Matsumoto, from the Cenomanian of Japan and Alaska, 
is a clear synonym of P. suhalpinum ellipticum, whilst the forms compared with this 
subspecies by Pervinquiere (1907, 1910), Crick (1907a) and Collignon (1928) clearly 
belong to it. The specimen of Phylloceras cf. ellipticum described by Collignon in 

1956 from the Turonian of Madagascar appears to be a P. {Hypophylloceras) 
improvisum Stoliczka ; the Phylloceras ellipticum of Parona & Bonarelli (1897) 
from the Albian of Escragnolles should be referred to P. [Hypophylloceras) aphrodite. 

P. (H.) suhalpinum ellipticum is a distinctive form. Whorl section alone readily 
separates it from P. (H.) suhalpinum suhalpinum, as noted above, whilst forms such 
as P. {H.) fortunei and P. {H.) sardoum have distinctly trigonal whorl sections. 
P. [H.) improvisum is a form with rounded whorls and a broad venter, and P. [H.) 
aptiense aptiense and applanatum and P. (H.) aphrodite all bear constrictions 
throughout ontogeny. P. (Hypophylloceras) thetys thetys, thetys diegoi and thetys 
majoricense, P. (H.) moreti and P. (H.) suhseresitense are as readily distinguished, 
being ornamented by fine striae rather than the low folds of P. (H.) suhalpinum 
ellipticum, and there are other obvious differences (Wiedmann 1964 ; Wiedmann & 
Dieni 1968). 

Occurrence. Phylloceras [Hypophylloceras) suhalpinum ellipticum is known 
from the Lower Albian of the Balearics, the Upper Albian of Sardinia, Tunisia, 
southern Russia and South Africa (Zululand), and the Cenomanian of Algeria, 
southern India, Madagascar, South Africa (Zululand), Alaska and Japan. 

Phylloceras (Hypophylloceras) aphrodite Fallot & Termier 
(PI. 3, fig. 3) 

1897 Phylloceras ellipticum Kossmat ; Parona & Bonarelli : 78 ; pi. 10, fig. 7. 

1923 Phylloceras aphrodite Fallot & Termier : 25 ; pi. 2, figs 5-7 ; text-fig. 5. 

1947 Phylloceras escragnollensis Breistroffer : 55. 

1955 Salfeldiella [Aphroditiceras) aphrodite var. venus Mahmoud : 75 ; pi. 5, fig. i ; text-fig. 43. 

1957 Phylloceras aphrodite Fallot & Termier ; Almela & Revilla : 19 ; pi. 3, fig. 5. 
1957 Phylloceras [Salfeldiella] guettardi Raspail ; Almela & Revilla : i6 ; pi. 2, fig. 6. 
1957 Phylloceras aft. grothi Fallot ; Almela & Revilla : 16 ; pi. 2, fig. 7. 

1962a Hypophylloceras aphrodite (Fallot & Termier) ; Wiedmann : 252 ; pi. 16, fig. 5 ; text- 
fig- 5- 

1962b Hypophylloceras aphrodite (Fallot & Termier) ; Wiedmann : 141. 

1964 Ph. [Hypophylloceras] aphrodite Fallot & Termier ; Wiedmann : 184 ; pi. 18, figs 4-6 ; 
pi. 20, fig. I ; text-fig. 40. 

1968 Ph. [Hypophylloceras] aphrodite Fallot & Termier ; Wiedmann & Dieni : 25. 

Neotype. Designated by Wiedmann, 1962a : 252 ; refigured by him in 1964 : 
pi. 18, figs 4a-c. From the Upper Albian of Son Vida-Son Suredeta, Mallorca 

Material. Two specimens. BM(NH) C78667 from Bed 6, Loc. 51 and 
SAS PJR/i from Loc. 57, both along the Mzinene River, Zululand (Albian V). 


Dimensions. D Wb Wh WbjWh U 

SASPJR/i 34-0 147(43) 17-0(50) 0-87 5-0(7) 

BM(NH) C78667 37-0 i6-9 (46) 20-0 (54) 0-84 5-0 (13) 

Description. The coiling is very involute, with a small, deep, conical umbilicus 
(7-13% of diameter). The whorl section is slightly compressed, suboval, with the 
greatest breadth at mid-flank. The umbilical wall is subvertical, sloping outwards ; 
the umbilical shoulder is abruptly rounded, the flanks convex and markedly inflated 
and the venter broadly and evenly rounded. The surface of the test bears fine 
flexuous striae, and irregular low folds which arise at the umbilical seam, sweep 
forwards across the inner flank, slightly backwards across mid-flank and forwards 
again across shoulder and venter. There are four to five deep, narrow, flexuous 
constrictions per whorl, paralleling the growth striae, which are weak at the 
umbilical shoulder but deep across the flank and venter. The internal mould is 
virtually smooth save for constrictions. 

The suture includes a rather narrow diphyllic E/L saddle, separated by a trifid 
lobe from a smaller but also diphyUic L/Ug. 

Discussion. The presence of constrictions throughout ontogeny serves to 
distinguish this species from most other members of the thetys group. P. {Hypo- 
phylloceras) aptiense aptiense is more compressed (whorl breadth to height ratio is 
typically 0-7) and has broader, straighter constrictions ; P. (H.) aptiense applanatum 
has an equally compressed whorl, flat, subparallel flanks and a somewhat flattened 
venter (Wiedmann 1964 : text-fig. 43a). 

Occurrence. Upper Albian of the Balearics, Sardinia, southern France 
(Escragnolles), Sinai (Moghara Massif) and South Africa (Zululand). 

2. Group of P. {Hypophylloceras) velledae (Michehn) 

This species group is transitional between the thetys and seresitense groups, having 
diphyllic saddles during early ontogeny and tetraphyllic saddles when adult. The 
following are referred to the group : 

1. Hypophylloceras velledae velledae (Michelin) (1834 • pl- 35) » Albian to Cenomanian. 

2. Hypophylloceras velledae (Michelin) morelianum (d'Orbigny) (1840 : 176 ; pi. 54, 
figs 1-3), Aptian to Albian. 

3. Hypophylloceras velledae (Michelin) aschiltaense Breistroffer (1947 : 55), Aptian 
to Albian. 

4. Hypophylloceras velledae (Michehn) inflatum CoUignon (1950 : 35 ; pi. 4, figs 
i6-i6b), Albian. 

5. Hypophylloceras ponticuli (Rousseau) (1842 : 783), Aptian. 

6. Hypophylloceras onoense Stanton 1895 [= Ammonites ramosus Gabb 1864 : 65 ; 
pi. II, fig. 12 ; non Meek], Aptian. 

7. Hypophylloceras cypris cypris Fallot & Termier (1923 : 23 ; pi. 3, figs la-c, 2c), 
Aptian to Albian. 

8. Hypophylloceras cypris Fallot & Termier cytherae Wiedmann (1964 : 218 ; pi. 12, 
figs 2a-c, 7a-b), Aptian to Albian. 


9. Hypophylloceras pervinquierei Collignon (1928 : 147, text-fig. 2 ; pi. XV, figs 9, 
9a), Lower Cenomanian. 

Phylloceras (Hypophylloceras) velledae (Michelin) 

This is the most commonly cited Albian phylloceratid, and the most frequently 
misinterpreted. Wiedmann (1964 : 205-207) has now clarified much of the con- 
fusion surrounding the species, and has designated a neotype for the restricted form. 
The species can be subdivided as follows : 

1. Whorl section compressed, oval, maximum whorl breadth at umbilical wall, 
sides flat - P. {H.) velledae morelianum (d'Orbigny), Aptian ? to Albian. 

2. Whorl section compressed, oval, sides moderately inflated, greatest breadth on 
inner third of flanks - P. (H.) velledae velledae (Michelin), Albian to Cenomanian, 

3. Whorl section compressed, elliptical, flanks inflated, greatest breadth at mid- 
flank - P. [H.) velledae aschiltaense Breistroffer, Aptian to Albian. 

4. Whorl section subcircular, maximum whorl breadth on inner third of strongly 
rounded flanks - P. {H.) velledae inflatum Collignon, Albian. 

Phylloceras (Hypophylloceras) velledae velledae (Michelin) 
(PI. 4, fig. 5 ; PI. 5, fig. 4 ; PI. 6, fig. I ; PI. 7, figs 2-3 ; Pis 8, 10, 11) 

1834 Ammonites velledae Michelin : pi. 35. 

1 84 1 Ammonites velledae Michelin ; d'Orbigny : 280 ; pi. 82, figs 1-4. 

1907b Phylloceras sp. Crick : 236. 

1921a Phylloceras velledae (Michelin) ; Spath : 273. 

1925 Phylloceras velledae (Michelin) ; Spath : 181 -182. 

1928 Phylloceras velledae (MicheHn) ; Collignon : 6 ; pi. i, figs i-ia. 

1930 Phylloceras velledae (Michelin) ; Besairie : 212 ; pi. 17, fig. 2. 

1936 Phylloceras velledae (Michelin) ; Venzo : 66 ; pi. 5, figs 4a-b. 

1947 Phylloceras velledae (Michelin) ; Breistroffer : 55. 

1949a Phylloceras velledae (Michelin), var. ; ColHgnon : 44 ; pi. 7, figs i-ia ; text-figs 4, 5. 

i960 Euphylloceras velledae Michelin ; Drushchitza & Kudryavtseva : 252 ; pi. 2, figs 5a-b. 

1962a Hypophylloceras velledae velledae (Michelin) ; Wiedmann : 250 ; pi. 16, fig. 4. 

1962b Hypophylloceras velledae velledae (Michelin) ; Wiedmann : 142. 

1963 Phylloceras velledae (Michelin) ; Collignon 14; pi. 241, figs 1036. 

1964 Phylloceras velledae (Michelin) ; Collignon 14; pi. 318, fig. 1347. 

1964 Phylloceras {Hypophylloceras) velledae velledae (Michelin); Wiedmann : 209 ; pi. 11, 
figs la-c ; pi. 13, figs 4a-b ; pi. 21, figs 4a-b ; text-fig. 49. (With full synonymy.) 

1968 Ph. [Hypophylloceras) velledae velledae (Michelin) ; Wiedmann & Dieni : 25 ; pi. i, 
figs 5a-b. 

Neotype. Designated by Wiedmann, 1964 : 211. MHNP 1895-17 (d'Orbigny 
Collection) from the Albian of Epothemont, near Bar-sur-Aube, Aube, France. 

Material. Numerous specimens. SAS LJE 86 from Loc. 163, Mfongosi 
Creek (Aptian III) ; BM(NH) C78685 from Loc. 36 on the Mzinene River (Albian 
III) ; BM(NH) C78668 from Bed i at Loc. 51, on the Mzinene (Albian IV) ; 
BM(NH) C78677, C78670 and C78683 from Bed 6, C78684, C78682 from Beds 12-13 


and BM(NH) C78669 (loose) also from Loc. 51 (Albian V) ; BM(NH) C78678, 
? C78679, SAS Z1503, Z1491, Zi492a-b, from Loc. 54 on the Mzinene ; BM(NH) 
C78671-6 from Loc. 56 on the Mzinene ; SAS A1847, from Loc. 57 on the Mzinene ; 
all Albian V. SAS A2053, PJR/2, Z1195, Z564, Z1248, Z1247 from Loc. 64 ; 
SAS 26/3214, 26/3227, 26/3223, BM(NH) C78680-1 and C78689 from Loc. 65, and 
SASA1189 from Loc. 66, all on the Munywana (Albian V). SAS Zi7(5), Zi7(6) 
and Zi9a-b come from Haughton's (1936 : 291 ; fig. 3) localities 17 and 19 in 
northern Zululand. 

The specimen referred to by Spath (1921a : 273) from the 'Middle Branch of the 
Manuan Creek', SAM 4992, also belongs to this species, as does Crick's (1907b) 
Phylloceras sp., BM(NH) C18264. 




Wh WblWh U 



61-0 (37) 

98-0 (59) 0-62 — 

Collignon 1949b : 45 


44-0 (37) 

69-0(59) 0-63 5 (4) 

CoUignon 1963 : 4 


43-0 (38) 

68-0 60) 0-63 3 (3) 

SAS 3214 


37-0 (37) 

57-2 (57) 0-65 6-0 (6) 

SAS Z19 


40-0 (35) 

70-0 (60) 0-57 5-5 (5) 



477 (35) 

83-2 (60) 0-57 6-8 (5) 

SAS Zi492a 


55-0 (34) 

97-2 (60) 0-57 6-5 (4) 

SAS Z1496 


6o-2 (36) 

103-2 (61) o-6o 9-5 (6) 

BM(NH) C78668 


77-5 (37) 

121-0 (58) 0-63 11-5 (6) 

Description. The 


is involute, 

with a tiny umbilicus, 3-7% of the 

diameter, with an outwards sloping wall. 

The whorl section is compressed, the 

whorl breadth to height ratio varying from 0-55 to 0-63, with the greatest breadth 
well below mid-flank. The sides are gently rounded, and converge from the lower 
flanks towards a quite broad, rounded venter. Ornament consists of fine flexuous 
striae, hair-like around the umbilicus, but coarsening and flattening on the upper 
flank and venter where they are approximately as wide as the interspaces ; there 
are about two hundred per whorl. They sweep gently forwards across the inner 
flank, backwards across mid-flank, and then forwards across the upper flank to pass 
across the venter with a shallow forward projection. There are no constrictions 
nor do the striae bunch. Internal moulds bear traces of the same ornament as the 
shell surface. 

The suture of large specimens includes an asymmetrically diphyllic saddle E/L, 
separated by a trifid lateral lobe from an L/U2 saddle, with five foholes (PI. 6, fig. 3a). 

Discussion. Wiedmann gives a complete synonymy for this form, and demon- 
strates very clearly how few of the specimens referred to Phylloceras velledae actually 
belong to the restricted species. Our collections contain, however, a large number 
of specimens ranging from juveniles to individuals which are still wholly septate at 
diameters of over 200 mm and match the neotype in proportions, style of ornament 
and suture. There is some variation in relative proportions and in strength of 
ornament, but all the material belongs to a single form. Our material appears to 
be the largest described collection of P. [H.) velledae velledae, and an ontogenetic 
series and a number of adult variants are figured here. 


Differences between P. [Hypophylloceras] velledae velledae, morelianum, aschiltaense 
and inflatum have already been given. The other Albian members of this group, 
P. [H.) cypris cypris and P. {H.) cypris cytherae, both show distinctly fasciculate 
ornament ; the former has a subtrigonal whorl section, and both have a more open 

P. {Hypophylloceras) seresitense, the other species of Hypophylloceras represented 
by large individuals in our collections, has a much wider umbilicus, and has much 
finer, straighter striae at similar diameters. 

Occurrence. Lower Albian of the Balearics ; Upper Albian of France ; Middle 
Aptian and Upper Albian of South Africa ; Upper Albian of Mozambique ; Lower 
Albian to Cenomanian of Madagascar. 

Phylloceras (Hypophylloceras) velledae (Michelin) inflatum Collignon 

(PI. 13, fig. I) 

1950 Phylloceras velledae Michelin var. inflata Collignon : 35 ; pi. 4, figs i6-i6b. 
1962a Phylloceras velledae inflatum Collignon ; Wiedmann : 142. 

1963 Phylloceras velledae Michelin var. inflata Collignon : 6 ; pi. 242, fig. 1040. 

1964 Phylloceras velledae inflatum Collignon ; Wiedmann : 207. 

Lectotype. The specimen figured by Collignon (1950) as his pi. 4, figs i6-i6b, 
from the Lower Albian of Komihevitra, Madagascar, is herein designated lectotype. 

Material. One specimen only, SAS 1742, from the Middle or Upper Albian of 
the Mzinene River, Zululand. 









49-0 (50) 

58-0 (59) 


2-0 (2) 

Colhgnon 1963 : 6 


52-0 (50) 

64-0 (62) 


4-0 (4) 

SAS 1742 


40-0 (50) 

45-3 (56) 


4-0? (5?) 

Description. The coiling is very involute, with a tiny, occluded umbilicus. 
An outward-sloping umbilical shoulder produces a conical circumbilical depression. 
The whorl section is subrounded, the whorl breadth to height ratio being 0-88. 
The whorl sides are inflated and the venter broadly rounded ; the greatest breadth 
is close to the umbilical shoulder. Ornament consists of fine, dense, flexuous 
striae, weak on the lower flank, but strong on the upper flanks and venter. The 
striae arise at the umbilical seam, pass straight across the lower flank, flex gently 
backwards at mid-flank, and gently forwards across shoulders and venter. The 
internal mould bears weaker but otherwise similar ornament. 

Discussion. The occluded umbilicus, whorl section and ornament clearly place 
this specimen in P. [Hypophylloceras) velledae inflatum ; it is identical with the 
Malagasy material figured by Collignon, save for slight differences in relative pro- 
portions. This form is readily distinguished from other subspecies of velledae, none 
of which approaches its degree of inflation. Form of umbilicus and ornament, as 
well as inflation, clearly differentiate it from other members of the velledae group. 


Occurrence. Lower Albian of Madagascar ; Middle or Upper Albian of South 
Africa (Zululand). 

Phylloceras (Hypophylloceras) sp. aff. velledae (Michelin) 

(PI. 5, fig. I ; PI. 7, fig. I) 

cf. 1972 Hypophylloceras aff. californicum (Anderson) ; McLearn : 22 ; pi. i, figs la-b, 2a-b. 

Material. Three specimens : BM(NH) C78696, from Loc. 51, collected loose 
but of Upper Albian age ; BM(NH) C78695 from Loc. 56 ; SAS A2239 from Loc. 57 ; 
all Mzinene River, Zululand (Albian V). 







BM(NH) C78695 


31-0 (38) 

46-5 (57) 


5-0 (6) 

BM(NH) C78696 


43-5 (40) 

64-2 (59) 


7-2 (6) 

SAS A2239 



89-5 (60) 


9-1 (6) 

Description. Three specimens in our collection appear to belong to the velledae 
group, but differ from P. {H.) velledae velledae in having a broader whorl section and 
from velledae velledae and velledae inflatum in their coarser ornament, effaced on the 
inner flank and not obviously fiexuous. McLearn has recently figured (1972 : pi. i, 
figs 1-2, especially 2) similar specimens from the Lower Albian of the Queen 
Charlotte Islands, British Columbia, referring to them as Hypophylloceras aff. 
californicum (Anderson). The type of this species is, however, a finely ornamented 
form, known only as a fragment from the Lower Albian of California (Anderson 
1938 : 143 ; pi. 12, fig. 7) and which is too poorly described and figured for adequate 

Occurrence. Upper Albian of Zululand. Lower and Upper Albian of the 
Queen Charlotte Islands, British Columbia. 

3. Group of P. {Hypophylloceras) seresitense Pervinquiere 

This group, characterized by possession of tetraphylloid saddles even in the early 
stages, includes seven Lower Cretaceous forms, most of which are listed by Collignon 
(1956 : 29 et seq.). Included here are the Neophylloceras, Epiphylloceras and 
Hyporbulites of previous workers. 

Phylloceras (Hypophylloceras) seresitense Pervinquiere 

The species can be divided as foUows (Wiedmann 1964 : 221). 

1. Whorl section moderately compressed, whorl breadth to height ratio up to 0-65, 
umbilicus narrow - P. (H.) seresitense seresitense Pervinquiere, Aptian to 

2. Whorl section very compressed, whorl breadth to height ratio 0-50-0-57, 
umbilicus narrow - P. {H.) seresitense boulei Collignon, Albian to Cenomanian. 

3. Whorl section high, umbilicus moderately open - P. {H.) seresitense tanit 
Pervinquiere, Albian to Cenomanian. 



Phylloceras (Hypophylloceras) seresitense seresitense Pervinquiere 
(PI. 4, fig. 6 ; PI. 6, fig. 4 ; PI. 7, fig. 4 ; PI. 9) 

1865 Ammonites velledae Michelin ; Stoliczka : 116 ; pi. 59, fig. i. 

1895 Phylloceras velledae Michelin ; Kossmat : 12 ; pi. i, fig. 3. 

1907 Phylloceras velledae var. seresitense Pervinquiere : 52. 

1907a Phylloceras velledae Michelin ; Crick : 166 ; pi. 10, figs lo-ii. 

191 o Phylloceras velledae var. seresitensis Pervinquiere ; Pervinquiere : 9, text-fig. 2 ; 

pi. I. figs 1-3. 
non 1923 Phylloceras seresitense Pervinquiere ; Spath : 18 ; pi. i, fig. 2 ; pi. 2, fig. i 

[= P. seresitense tanit Pervinquiere]. 
? 1928 Phylloceras ex aff. ramosi Meek ; Collignon : i ; pi. i, figs 2-4. 
? 1936 Phylloceras seresitense PervinquieTe ; Collignon : 190 ; pi. 21, figs 1-2. 
1940 Phylloceras seresitense Pervinquiere ; Fabre : 211 ; pi. 5, fig. i. 
1943 Phylloceras velledae Michelin ; Matsumoto : 676. 
1947 Hyporbulites seresitensis Pervinquiere ; Breistroffer : 82. 
1950 Phylloceras [Hyporhulties) seresitense Pervinquiere ; Collignon : 66. 
1956 Hyporbulites seresitensis var. raynaudiensis Collignon : 16 ; pi. 4, fig. i. 
1959 Neophylloceras seresitense (Pervinquiere) ; Matsumoto : 55, text-fig. 3 ; pi. 12, 

figs 4-5. 
1962a Hypophylloceras seresitense seresitense (Pervinquiere) ; Wiedmann : 142, text-fig. 

8 ; pi. 8, figs 1-2. 
1962b Hypophylloceras seresitense seresitense (Pervinquiere) ; Wiedmann : 249 ; pi. 16, 

fig. I. 

1963 Phylloceras [Hyporbulites) seresitensis Pervinquiere ; Collignon : 4 ; pi. 241, fig. 
1038 ; pi. 142, fig. 1041. 

1964 Ph. {Hypophylloceras) seresitense seresitense Pervinquiere ; Wiedmann : 221 ; pi. 15, 
figs 4a-b ; pi. 21, figs la-b ; text-fig. 52. 

1968 Phylloceras [Hypophylloceras) seresitense seresitense Pervinquiere ; Wiedmann & 
Dieni : 26. 

Lectotype. The original of Pervinquiere 1910 : pi. i, fig. i, designated by 
Wiedmann (1964). 

Material. Four specimens : BM(NH) C78690, C18137-8 from the Skoenberg 
(? Cenomanian III), and SAS A1401 from Loc. 36 on the Mzinene River (Albian 
III), all from Zululand. 

Dimensions. D Wb 

BM(NH) C18138 28-3 10-6 (37) 

SAS A1401 190-0 69-3 (36) 

Description. Our specimens add nothing to knowledge of the early develop- 
ment of this species, which is well known. We have, however, a near adult specimen 
(SAS A1401, PI. 9) still wholly septate at 190 mm, which appears to be the largest 
known individual referred to it. The coiling is very involute, the umbilicus small, 
conical and deep (5% of diameter) with a rounded undercut wall and an outwards 
sloping shoulder which merges with the flanks. The whorl section is compressed, 
the greatest breadth being one-third of the way up the flanks. The sides are gently 
inflated, converging to a broad rounded venter. 

Ornament consists of dense, fine striae, flat-topped, and as wide as the interspaces. 
There are between 400 and 450 of these on the outer whorl. They arise at the 




167 (59) 


1-5 (5) 

109-2 (57) 


10-5 (5) 


umbilical seam and are very faint, but strengthen across the flanks. They flex 
gently forwards on the inner flank, backwards over the outer flank and forwards 
across the shoulders to pass straight across the venter. The major saddles of the 
suture (E/L and L/U2) are tetraphylloid. 

Discussion. Our smaller specimens, originally figured by Crick as Phylloceras 
velledae Stoliczka (? Michelin), clearly belong to P. {H.) seresitense seresitense, as 
indicated by Wiedmann (1964). Our adult specimen, still wholly septate at a 
diameter of 190 mm, is referred to P. (Hypophylloceras) seresitense on the basis of 
the style and strength of ornament, relative proportions and such of the suture line 
as is visible. The whorl breadth to height ratio clearly excludes reference to 
P. {H.) seresitense boulei, but separation from P. {H.) seresitense tanit cannot be 
definitely confirmed at this size because the largest described tanit has a diameter 
of only 62 mm. It seems, however, that the umbilicus of tanit is usually 10% or 
more of the diameter. 

Occurrence. P. [Hypophylloceras) seresitense seresitense ranges from the Upper 
Aptian (Balearics) to the Cenomanian. The geographic distribution includes 
southern France, northern Spain, north Africa, the Balearics, southern India, 
Madagascar, South Africa and Alaska. 

Phylloceras (Hypophylloceras) androiavense Besairie 

(PI- 4, %• 7) 
1936 Phylloceras androiavensis Besairie : 165 ; pi. 16, figs 9-11. 
1963 Phylloceras {Hyporbulites) androiavense Besairie ; Collignon : 4 ; pi. 241, fig. 1037. 

Lectotype. Besairie's figured specimen (1936 : pi. 16, figs 9-11), from the 
upper part of the Middle Albian of Androiavy, Madagascar, is herein designated 

Material. One specimen only, BM(NH) C78667, from Loc. 54, Mzinene River, 
Zululand (Albian V). 

Description. Our specimen is fragmentary, with an original diameter of 
approximately 40 mm. The coiling is very involute, with a tiny umbilicus (approxi- 
mately 5% of the diameter). The whorl section is very compressed (whorl breadth 
to height ratio about 0-46), the greatest breadth being at the umbilical shoulder. 
The whorls are rapidly expanding, the whorl height being approximately 60% of 
the total diameter. The umbilical shoulder is quite abruptly rounded, the sides 
flat and subparallel with a narrow rounded venter. Ornament consists of fine, 
flexuous striae grouped into bunches ; about ten low, fold-like ribs arise at the 
umbilical shoulder, pass forwards across the inner flank, and flex backwards at mid- 
flank where they give rise to numerous striae which sweep gently forwards across 
the shoulder and venter, together with similar intercalated striae. The inner 
flanks between folds bear fine irregular striae. The sutures are not exposed. 

Discussion. Our specimen is identical in style of ornament, whorl section and 
(as far as can be determined) relative proportions to P. {Hypophylloceras) androia- 
vense. Apart from this species, fasciculate ornament is present in the following 




mid-Cretaceous forms. P. {Hypophylloceras) cypris cypris Fallot & Termier (1923 : 22 ; 
pi. 3, figs la-c, 2c) is an Albian form, differing, however, in having a subtrigonal 
whorl section, a whorl breadth to height ratio of 0-66, and a larger umbilicus 12% 
of the diameter. P. {Hypophylloceras) cypris cytherae Wiedmann (1964 : 218 ; 
pi. 12, figs 2a-b, 7a-b ; pi. 15, fig. 8a-b ; text-fig. 12) is an Upper Aptian to Albian 
form with a compressed, oval whorl section, rounded flanks, a whorl breadth to 
height ratio of 0-66 and a larger umbilicus 12% of the diameter. Hypophylloceras 
algeriense Wiedmann (1962 : 144 ; pi. 8, fig. 3a-b ; text-fig. 9), an Albian to 
Cenomanian species with a whorl breadth to height ratio of 0-65, has rounded, 
convergent flanks and an arched venter. 

Occurrence. High Middle Albian of Madagascar ; Upper Albian of South 
Africa (Zululand). 

Phylloceras (Hypophylloceras) woodsi woodsi van Hoepen 
(Figs 4-6 ; PI. 13, figs 3-5) 

1906 Phylloceras sp., Woods : 331 ; pi. 41, fig. 4. 

1 92 1 Phylloceras woodsi van Hoepen : 3 ; pi. 2, figs i 
1921b Phylloceras nera (Forbes) ; Spath : 40. 

1922 Phylloceras woodsi van Hoepen ; Spath : 117. 

1929 Phylloceras woodsi van Hoepen ; Barrabe : 174 ; pi. 

1930 Phylloceras woodsi van Hoepen ; Besairie : 567 (223) 

1931 Phylloceras woodsi van Hoepen ; Collignon : 10. 
1956 Hyporhulites woodsi van Hoepen ; Collignon : 19. 
1956 Hyporhulites hoepeni Collignon : 19 ; pi. i, figs 4, 4b, 5, 5b, 
1966 Phylloceras {Hyporhulites) hoepeni Collignon ; Collignon : i 

text-fig. I. 

fig. 17. 

pi. 21, figs 2- 

6-6b ; text-fig. 4. 
: pi. 455, fig. 1406. 

HoLOTYPE. TM 537, figured by van Hoepen (1921 : pi. 2, figs 3-4). 


4. External suture line of Phylloceras (Hypophylloceras) woodsi woodsi van Hoepen. 
Holotype, TM 537, x 12-5. 


Fig. 5. Partial external suture of Phyllo- 

ceras [Hypophylloceras) woodsi woodsi van 

Hoepen. TM 529, x 6. 

Material. Eight specimens : the holotype TM 537 ; four paratypes, TM 529, 
530, 533, 534 ; a specimen in the Durban Museum (unregistered) ; a specimen in 
the Griesbach Collection, jfigured by Woods (1906) ; and BM(NH) C78826. All are 
from the Umzamba Formation of Loc. i, Umzamba, Pondoland, and are of Santonian 
or Campanian age. 

Description. The coiling is moderately involute (umbilicus approximately 
7-10% of diameter). The whorl section is a compressed oval with virtually parallel 
flanks, converging only slightly towards the gently rounded venter. Maximum 
whorl width is at the middle to inner third of the flanks. The umbilicus is pit-like, 
with nearly vertical walls, thus forming a pronounced, angular umbilical shoulder. 

Ornament consists of numerous, fine single ribs which are most pronounced on 
the outer third of the flanks. Ribbing on the inner third is very faint, but the ribs 
appear to originate at the umbilical edge, flex slightly backward on the inner third 
of the flanks and gently forward at mid-flank, and then pass in a prorsiradiate 
direction across the ventrolateral shoulders. 

The suture is typically phylloid on the inner whorls, but tends to become more 
dendritic at the largest diameters studied. E/L is diphylloid, separated from the 
triphylloid L/Ug by a trifid lateral lobe. 

Discussion. CoUignon (1956 : 19), when erecting P. {H.) hoepeni, stated that 
the species was very close to P. [H.) woodsi ; as far as relative proportions are 
concerned, the two species overlap to a great extent. P. (H.) hoepeni has a smaller 
umbilicus than the present species, 3-6% of diameter as against 7-10% in P. {H.) 
woodsi, but CoUignon realized that this criterion is difficult and rather impracticable 
to apply. He stated, in addition, that the suture lines were different ; in P. {H.) 
hoepeni the suture line is more zigzag-like and the saddles have smaller stems and 
are more incised. Unfortunately, only E/L and part of the lateral lobe of P. [H.) 
hoepeni have been figured by CoUignon (reproduced here as Fig. 6), whilst the 
suture line given by van Hoepen for P. (H.) woodsi is rather schematic. 



Fig. 6. Partial external suture line of 
' Hyporbulites' hoepeni Collignon (= P. 
{Hypophylloceras) woodsi woodsi van Hoepen 
herein), after Collignon (1956 : text-fig. 4), 

Virtually the same part of the suture line as figured by Collignon is visible on one 
of the paratypes of P. (H.) woodsi, TM 529 (Fig. 5) ; it is quite clear that, apart 
from the folioles in the saddles of P. {H.) hoepeni being more phylloid than in 
P. {H.) woodsi, the two suture lines are identical. Because of the comparable 
relative proportions, similar suture lines and (probably) the same stratigraphic 
occurrence, P. {H.) hoepeni is here regarded as a junior synonym of P. [H.) woodsi. 
P. {H.) hoepeni infundibuliformis Collignon is also referred to P. {H.) woodsi as a 
subspecies. It is more inflated than the nominal subspecies and stands in the same 
relationship to P. [H.) woodsi that P. [H.) suhalpinum ellipticum does to P. (H.) 
subalpinum suhalpinum. 

Less readily resolved is the relationship of P. {H.) woodsi to 'Neophylloceras' 
marshalli Shimizu [= Phylloceras nera Marshall (non Forbes) 1926: 124; pi. 19, 
fig. 4 ; pi. 26, figs 1-2], 'N.' hetonaiense Matsumoto (1943 : 675, text-figs lag, bg) of 
which 'N.' lamhertense Usher (1952 : 50 ; pi. i, figs 1-3) is a synonym (Jones 1963 : 
23), and 'N.' nera Forbes (1846 : 106 ; pi. 8, fig. 7). All have ornament well within 
a range of intraspecific variation which we would find acceptable in phyUoceratids. 
In the suture line of N. hetonaiense, however, the elements of the suspensive lobes 
form a concave arc ; in N. nera they lie on a straight line, and in the present species 
the suspensive lobe is slightly retracted (Fig. 4) . Given better collections, it may be 
possible to show that all these species are junior subjective synonyms of P. {Hypo- 
phylloceras) nera. 

Occurrence. Santonian or Campanian of Zululand ; Santonian Texanites 
oliveti Zone, Madagascar. 

Phylloceras (Hypophylloceras) tnikobokense (Collignon) 

(PL 12, fig. I) 

1938 Phylloceras aff. surya (Forbes) ; Collignon : 41 ; pi. 7, fig. i. 

1956 Epiphylloceras mikobokense Collignon : 24 ; pi. 2, figs 3-3a ; pi. 4, figs 5 -5b. 

HoLOTYPE. MNHP 1567, figured by Collignon (1956 : pi. 4, figs 5-5b), from the 
Lower Maastrichtian of Lajona, Madagascar. 

Material. Two specimens, SAS H163C/7 and 163/9, from Loc. 20, Mfolozi 
River, Zululand (Maastrichtian I). 


Dimensions. (From CoUignon 1956 : 24) 

D Wh Wh WblWh U 

EMP 1567 86-0 25-0 (29) 42-0 (49) 59 — 

EMP 1564 73-0 23-0 (32) 42-0 (58) 55 ii-o (15) 

Description. Our specimens are fragments of discoidal shells with a moderately- 
high expansion rate and a compressed whorl section (whorl breadth to height ratio 
is 0-51), the greatest breadth being close to the umbilical shoulder. The umbilicus 
is small, about 10% of the inferred diameter, quite deep, with a subvertical, rounded 
wall and an abruptly rounded shoulder. The flanks are rounded, converging to a 
narrowly rounded venter. 

Ornament consists of fasciculate ribs and striae. Our specimens bear strong 
fold-like ribs which arise at the umbilical seam, pass straight across the lower flank, 
thicken and flex backwards across the outer flank, and die out across the ventro- 
lateral shoulder, being represented on the venter merely by low, broad swellings. 
Superimposed on these are fine, dense striae, arising at the umbilical seam, branching 
on the flanks and following a course parallel to the ribs. 

The suture is deeply incised, having lost the typical phylloid terminations 
(PI. 14, fig. lb). 

Discussion. The very coarse, fold-like ribs and fine striae, and relative pro- 
portions, indicate that our specimens belong to CoUignon's species, comparing 
closely with the large paratype fragment figured by him (1956 : pi. 2, figs 3-3a). 
Ornament readily separates this species from most Upper CretdiceousHypophyllocer as, 
whilst those forms with comparable fasciculate ornament {'Epiphylloceras' of 
authors) differ as follows. Hypophylloceras surya (Forbes) (1846 : 106 ; pi. 7, 
fig. 10) has much coarser ribbing rather than striae ; Hypophylloceras cottreaui 
CoUignon (1956 : 25 ; pi. 4, figs 6a-b) lacks strong fold-like ribs, having instead 
fasciculate striae arising from umbilical bullae. Epiphylloceras radiatum Marshall 
(1926 : 135 ; pi. 19, fig. 7 ; pi. 26, figs 3-4 ; see also Henderson 1970 : 6, text-fig. 
26) has a smaller umbilicus and less compressed whorls, the whorl breadth to height 
ratio being 072. 

Occurrence. Lower Maastrichtian of Madagascar and South Africa (Zululand). 

Genus PARTSCHICERAS Fucini 1920 

Type species. Ammonites partschi Stur 1851 (non Klipstein 1843), by the 
subsequent designation of Spath, 1927. 

Discussion. This genus is reviewed at length by Wiedmann (1962b : 257 ; 
1964 : 229). Our material is too poor to allow profitable discussion, and we have 
therefore followed him in regarding Phyllopachyceras Spath 1925, Procliviceras 
Fucini 1920, Macrophylloceras Spath 1927, Hoplophylloceras Spath 1927, Partchi- 
phylloceras Roman 1938 and ? Calaiceras Kovacs 1939, as synonyms. 

Occurrence. Sinemurian to Maastrichtian ; in the Cretaceous species have a 
world-wide distribution save for the boreal region of north-west Europe and the 
western interior of North America. 


Partschiceras umzambiense (van Hoepen) 
(Fig. 7 ; PI. 13, fig. 2) 

1920 Phylloceras umzambiense van Hoepen : 142 ; pi. 24, figs 1-3. 

1921b Phylloceras umzambiense van Hoepen ; Spath : 50. 

1922 Phylloceras umzambiense van Hoepen ; Spath : 117. 

1956 Phyllopachyceras umzambiense (van Hoepen) ; Collignon : 27 ; pi. 2, figs 2 -2b ; text-fig. 8. 

1966 Phyllopachyceras umzambiense (van Hoepen) ; Collignon : pi. 455, fig. 1850. 

HoLOTYPE. TM 524, figured by van Hoepen (1920 : pi. 24, figs 1-3), from the 
Umzamba Formation, Loc. i, Pondoland, and of Santonian-Campanian age. 
Material. The holotype only. 









17-0 (39) 

27-0 (61) 


2-0 (5) 

EMP 1587 

(Collignon 1956) 


lo-o (45) 

13-0 (59) 


i-o (5) 

Collignon 1966 : 

pi. 455, fig. 1850 


lo-o (49) 

12 (57) 


i-o (9) 

Description. The holotype is an incomplete, septate internal mould, badly 
abraded on one side. Coiling is involute with a small umbilicus, 5% of the diameter. 
The umbihcus is funnel-shaped and not distinctly demarcated from the flanks. 
Whorl section is ovoid with rounded flanks and a broad venter. Maximum breadth 
is at the inner third of the whorl. 

Ornament consists of thin, sinusoidal ribs, separated by somewhat wider inter- 
spaces. The ribs flex forwards on the inner half of the flanks and then backwards on 
the outer half to traverse the venter radially. Approximately eight pairs of fascicu- 
late ribs arise at the umbilicus as low, radial swellings which are confined to the 
umbilicus, disappearing on the flanks. Apart from the normal ribs, intercalatories 
or bifurcating ribs originate at about mid-flank. 

The main saddles (E/L and L/U2) are tetraphyllic. L is larger than Ug (Fig. 7). 

Discussion. This is a relatively rare species and has only been recorded on 
three occasions to date (van Hoepen 1921 ; Collignon 1956, 1966). The main 
distinguishing feature is the ornamentation of fasciculate ribbing near the umbihcus 
and intercalatory bifurcating ribbing on the vental part of the flanks. 

Partschiceras hesairiei hesairiei Collignon and P. hesairiei tsianalokensis (Collignon 
1931 : 10 ; pi. I, figs 2-2b, 3-3a, 4-4b ; pi. 8, fig. i) from the Santonian of 
Madagascar bear a superficial resemblance as far as inflation of the whorls is con- 
cerned, but has different ornamentation ; P. hesairiei hesairiei has strong ribs on 
the ventral area and P. hesairiei tsianalokensis lacks ribbing altogether. Of other 
Upper Cretaceous species, Parschiceras forbesianum (d'Orbigny 1850 : 213) (of which 
according to Henderson (1970 : 7) P. ezoense Yokoyama (1890 : 170 ; pi. 19, figs 
2a-c), P. minimum Marshall (1926 : 137 ; pi. 19, fig. 8 ; pi. 26, figs 5-6), P. histriatum 
Marshall (1926 : 138 ; pi. 19, fig. 5 ; pi. 27, figs 1-2), P. inflatum Shimizu (1935 : 
178), P. marshalli Collignon (1937 : 26) and P. zelandicum Collignon (1956 : 31) are 
synonyms), P. whiteavesi Kossmat (1897 : 124-125 [= Phylloceras forbesianum 


Fig. 7. External suture line of Partschiceras umzambiense (van Hoepen). Holotype, 

TM524, x6. 

Kossmat (non d'Orbigny) 1895 : 109 ; pi. 15, figs la-c ; = Ammonites rouyanus 
Stoliczka (non d'Orbigny) 1865 : 117 ; pi. 69, figs 5-7]) and P. occlusitm Tate 
(1865 : 36 ; pi. 3, figs la-b) all lack the fine, dense, wiry ribbing of P. umzambiense 
and are virtually smooth when preserved as internal moulds. 'Epiphylloceras' 
species, which may bear similar fasciculate ribbing, all differ from P. umzambiense 
in their compressed whorl section. 

Occurrence. Santonian of Madagascar [Texanites oliveti Zone) ; Santonian or 
Lower Campanian of Pondoland. 

Partschiceras sp. juv. ? cf. baborense (Coquand) 

(PI. 6, figs 2-3) 

cf. 1880 Ammonites baborensis Coquand : 26. 

cf. 1964 Partschiceras baborense (Coquand) ; Wiedmann : 243 ; pi. 14, figs 2, 4, 5 ; pi. 16, 
figs 1-2 ; pi. 21, figs 5, 6 ; text-fig. 59. (With synonymy.) 

Material. Three specimens, BM(NH) C78692-4, from Loc. 170, Mlambong- 
wenya Spruit, northern Zululand (Barremian I). 

Dimensions. D Wb Wh U 

BM(NH) C78693 i8-o — (— ) lo-i (56) — 

Description. The shell is involute, with an occluded umbilicus. The whorls 
are slightly compressed, the greatest breadth being close to mid-flank. Although 
the umbilicus is occluded, the umbilical wall slopes outwards to produce a shallow 
conical circumbilical pit before merging with the broadly rounded, subparallel 
flanks. The ventrolateral shoulder and venter are broadly and evenly rounded. 
The shell surface is quite without ornament, although this may be a result of partial 

The suture is too poorly exposed and corroded for detailed description. 

Discussion. These specimens are too poor for satisfactory determination. 
The nature of the umbilicus, lack of ornament and relative proportions recall. 


however, Partschiceras bahorense, especially pyritic juveniles figured by Wiedmann 
(1964) and Collignon (1937 : pi. i, figs 4-4b to 6-6b). 

Occurrence. P. bahorense ranges from Barremian to Aptian, and occurs widely 
in southern France, northern Spain, the Balearics, Sardinia, central Europe, the 
Caucasus, Bulgaria, the Crimea, north Africa and Madagascar. 

Genus CARINOPHYLLOCERAS KHnger, Wiedmann & Kennedy 1975 

Type species. Carinophylloceras collignoni Klinger, Wiedmann & Kennedy 
1975 : 658 ; pis 76-77 ; text-figs 1-3. 

Diagnosis. Phylloceratid ammonites with fastigate to distinctly keeled venters. 
Whorl section ovoid, higher than wide, with maximum width at umbilical margin ; 
narrowly umbilicated. Ornament typically phylloceratid, consisting of striae. 
Suture phylloid, with lituid I, trifid L, saddles E/L assymmetrically diphylHc, 
L/U2 asymmetrically tetraphyllic. Saddles in U3 asymmetrically diphyllic (Figs 

Discussion. Carinophylloceras is unique amongst Cretaceous phylloceratids by 
virtue of the presence of a keel. Keeled phylloceratids occur in the Jurassic, 
e.g. Harpophylloceras Spath 1927 and Menegheniceras Hyatt 1900. There are, 
however, no Cretaceous taxa referable to these genera and any affinity of Carino- 
phylloceras with these forms may be ruled out. 

The suture line of Carinophylloceras with an asymmetrical diphyllic saddle E/L 
and asymmetrical tetraphyllic L/U2, the ornamentation, degree of evolution and to 
a lesser extent whorl section point to affinities with the Albian/Cenomanian Phyllo- 
ceras {Hypophylloceras) velledae (sensu Wiedmann 1964), and to the Albian/ Aptian 
P. {H.) cypris cypris Fallot & Termier (Wiedmann 1964 : fig. 50 ; pi. 13, fig. 3 etc.). 
Apart from the keel, the whorl section is somewhat intermediate between P. {H.) 
velledae velledae and P. [H.) velledae morelianum. The presence of a keel, however, 
clearly separates Carinophylloceras collignoni from the group of P. {H.) velledae. 
Carinophylloceras is a homeomorph of Damesites, and was originally believed to be 
a new species of that genus. The similarities are remarkable. Not only the whorl 
section and the presence and shape of the keel, but also the degree of shell involution, 

Fig. 8. External suture line of Carinophylloceras collignoni Klinger, Wiedmann & 
Kennedy. UPE B464, x 1-3. 


the course of the ornamentation and even the external suture Hne show such 
similarities that these genera can scarcely be distinguished on their external 
morphology. Examination of the suture line, however, especially the internal part, 
reveals that it is typically phylloid with a lituid internal lobe (Fig. 9), a characteristic 
of all phylloceratinids (Wiedmann 1972 : 115). The internal lobe of Damesites as 
figured by Matsumoto (1954 : fig. 11) is intensively frilled and of desmoceratid type ; 
indeed no known desmoceratids possess a lituid internal lobe. 
Occurrence. Lower Albian (Albian III) of Zululand. 

Carinophylloceras collignoni Klinger, Wiedmann & Kennedy 

(Figs 8-9 ; PI. 14 ; PI. 15, figs 1-2) 

1975 Damesites ? sp. nov. Kennedy & Klinger : 276. 

1975 Carinophylloceras collignoni Klinger, Wiedmann & Kennedy : 658 ; pi. 76, figs la-b ; 
pi. 77, figs 1-3 ; text-figs 1-3. 

HoLOTYPE. SAS A1577, from the Mzinene Formation, stream cliff section along 
the Mzinene River 1200 m NE of the farm Amatis, north of Hluhluwe, Zululand, 
South Africa, 27°58'o3" S, 32°i8'34" E. Loc. 35 of Kennedy & Klinger (1975). 

Material. In addition to the holotype we have 39 paratypes : SAS UMS/2, 
SAS A1133 and BM(NH) C78639, C78644, C78647-8, C78767, C78769, C78770 from 
Loc. 35, and BM(NH) C78640-3, C78645-6, C78651 and C78768 from Loc. 36, 
both on the Mzinene River. SAS H93D/1, SAS H93/1, SAS H93/2, SAS 93/3, 
SAS H93/5, from Loc. 142, Nxala Estate, southern part of Mkuze Game Reserve, 
Zululand. SAS EM 91, SAS EM 92, SAS EM 77 from the Msunduzi Pan at 
26°57'25" S, 32°i2'4o" E ; UPE B 33 from the same area, at 26°57'io" S, 
32°i2'45" E. SAS EM 245a, b, c, SAS EM 93, SAS EM 244, SAS EM 114, from the 
Ndumu region, northern Zululand at 26°55'55" S, 32°i2'55" E. SAS LJE 134A, 
UPE B463, UPE B464, UPE B411 and BM(NH) C78649-50, from Loc. 174, and 
BM(NH) C78766 and C78771, from Loc. 171, both Mlambongwenya Spruit, northern 
Zululand. UPE B23, from Aloe Flats Estate, northern Zululand, at 26°59'5o" S, 
32°ii'5o" E. 

All specimens are from the Mzinene Formation of late early Albian age (Albian 







Holotype SAS A1577 


6o-5 (41) 

88-5 (59) 


8-5 (6) 

Paratypes SAS EM 244 


44-5 (36) 

68-0 (55) 


8-0 (6-5) 

SAS H98/1 


32-5 (41) 

44-5 (58) 


6-0 (8) 

SAS H93/3 


40-5 (37) 

63-5 (58) 


7-5 (7) 



47-0 (36) 

77-5 (58) 


9-0 (6-8) 

Description. Coiling is moderately involute with a narrow funnel-shaped 
umbilicus (6-8% of diameter). Whorl section is subtrigonal with a fastigate to 
distinctly keeled venter. Maximum width is at the umbihcal edge. In juvenile 
stages the venter is fastigate, but in the adult a distinct keel is developed. The 


Fig. 9. Partial internal suture line of 

Carinophylloceras collignoni Klinger, Wied- 

mann & Kennedy, showing overlapping 

lituid internal lobe. SAS EM 114, x 8-3. 

keel is of the floored type, and, depending on the mode of preservation, may be 
either present or absent on internal moulds. 

Ornament consists of pronounced striae which arise at the umbilical wall, are 
bent forwards at first, then sweeping gently backwards near the middle of the flanks, 
being finally strongly projected on the outer part of the flanks. They are bundled 
at their origin, and much stronger on the outer part of the whorls and venter, 
producing a chevron-like ventrolateral and ventral ornament. On internal moulds 
the ornamentation is still present, though very much subdued. 

Suture line as for genus. Auxiliary saddles in U3 are triphyllic. 

Occurrence. Lower Albian (Albian III) of Zululand. 


We are grateful to Dr H. W. Ball, Dr M. K. Howarth and Mr D. Phillips of the 
Department of Palaeontology, British Museum (Natural History), General 
M. Collignon (Moirans), Mr C. W. Wright (London), Professor Dr J. Wiedmann 
(Tubingen), Professor D. L. Visser (Pretoria), Dr C. K. Brain and Mrs E. Voigt 
(Transvaal Museum, Pretoria), Mr P. J. Rossouw (Pretoria), Mr M. R. Cooper 
(South African Museum, Cape Town) and the technical staff of the Department of 
Geology, Oxford, Geological Survey of South Africa, Pretoria, Institut fiir Geologic 
und Palaontologie, Tubingen, and Transvaal Museum, Pretoria, for their help and 
assistance in many ways. Thanks are also due to the Director, Geological Survey, 
Pretoria, for permission for one of us (H. C. K.) to publish these results. 


Almela, a. & DE LA Revila, J. 1957- Fosiles piritosos de Cretaceo de la Sierra de Ricote 
(Murcia). Boln Inst. geol. min. Esp., Madrid, 68 : 45-83, pis i-ii. 

Anderson, F. M. 1938. Lower Cretaceous deposits in California and Oregon. Spec. Pap. 
geol. Soc. Am., New York, 16, 244 pp., 84 pis. 

Arnould-Saget, S. 195 1. Les ammonites pyriteuses du Tithonique superieur et du Berriasien 
de Tunisie central. Annls Mines Giol., Tunis, 10 : 1-132, pis i-ii. 


Barrabe, L. 1929. Contribution a I'etude stratigraphique et petrographique du pays de 
Sakalave, Madagascar. M6m. Soc. geol. Fr., Paris, n.s. 12 : 1-270, pis i-io. 

Besairie, H. 1930. Recherches geologiques a Madagascar. Contribution a I'etude des 
ressources minerales. Bull. Soc. Hist. nat. Toulouse, 60 : 345-616 (1-272), pis 1-27. 

1936. Recherches geologiques a Madagascar. Premiere suite. La geologic du nord- 

ouest. Mem. Acad, malgache, Tananarive, 21 : 9-259, pis 1-24. 

Beznosov, N. V. 1958. Jurassic ammonites of the northern Caucasus and Crimea. Phyllo- 

ceratina and Lytoceratina. 118 pp, 34 pis. Gostoptekhizdat, Leningrad. [In Russian.] 
BouLE, M., Lemoine, p. & Thevenin, a. 1906-07. Paleontologie de Madagascar. Ill - 

Cephalopodes cretaces des environs de Diego-Suarez. Annls PaUont., Paris, 1 : 173-192, 

pis 14-20 (1906) ; 2 : 1-56, pis 1-8 (1907). 
Breistroffer, M. 1947- Sur les zones d'ammonites dans I'Albien de France et d'Angleterre. 

Trav. Lab. Geol. Univ. Grenoble, 26 : 1-88. 
BusNARDO, R. & David, L. i957- Contribution a I'etude des faunes d'Ammonoides de 

Medjez Sfa (Est Constantinois) . Pubis Serv. Carte geol. Alger., Algiers, n.s. 13 : 67-123, 

pis 1-3. 
CoLLiGNON, M. 1928-29. Les Cephalopodes du Cenomanien pyriteux de Diego-Suarez. 

Annls Paleont., Paris, 17:139-160 (1-24), pis 15-19 (1-5) (1928); 18:1-56 (25-79), 

pis 1-2 (6-7) (1929). 

1931- Faunes senoniennes du nord et de I'ouest de Madagacar. Annls geol. Serv. Mines 

Madagascar, Tananarive, 1 : 1-66, pis 1-9. 

1936. Fossiles du Turonien superieur d'Antanitiloky. In Besairie, H. Recherches 

geologiques a Madagascar. Premiere suite. La geologic du Nord-Ouest. M4m. Acad, 
malgache, Tananarive, 21 : 200-202, pi. 21. 

1937- Les ammonites pyriteuses de I'Aptien d'Antanatanamirafy. Annls Paleont., 

Paris, 26 : 107-132, pis 16-18. 

1938. Ammonites Campaniennes et Maastrichtiennes de I'ouest et du sud de Madagascar. 

Annls geol. Serv. Mines Madagascar, Tananarive, 9 : 55-118, pis 1-9. 

1949a. Faune neocomienne des couches a Crioceras de Belohasifaka (cercle de Sitampiky). 

Annls geol. Serv. Mines Madagascar, Tananarive, 15 : 53-83, pis 8-13. 

1949b. La faune albiene d'Ambarimaninga. Annls geol. Serv. Mines Madagascar, 

Tananarive, 16 : 1-128, pis 1-22. 

1950. L'Albien de Mokaraha. Annls geol. Serv. Mines Madagascar, Tananarive, 

17 : 57-85. pis 10-14. 

1956. Ammonites neocretacees du Menabe (Madagascar). IV. - Les Phylloceratidae. 

V. - Les Gaudryceratidae. VI. - Les Tetragonitidae. Annls geol. Serv. Mines Madagascar, 
Tananarive, 23 : 1-106, pis i-ii. 

1962. Atlas des fossiles caracteristiques de Madagascar {Ammonites). VIII {Berriasien, 

Valanginien, Hauterivien, Barremien) : 96 pp, pis 176-214. Service Geologique, Tananarive. 

1963. Atlas des fossiles caracteristiques de Madagascar {Ammonites). X {Albien) : 

XV+184 pp, pis 241-317. Service Geologique, Tananarive. 

1964. Atlas des fossiles caracteristiques de Madagascar {Ammonites). XI {Cenomanien) : 

xi-l-152 pp, pis 318-375. Service Geologique, Tananarive. 

1966. Atlas des fossiles caracteristiques de Madagascar {Ammonites). XIV {Santonien) : 

x-t-134 pp, pis 455-513. Service Geologique, Tananarive. 

Coquand, H. 1880. Etudes supplementaires sur la paleontologie algerienne. Bull. Acad. 

Hippone, Bone, 15 : 1-449. 
Crick, G. C. 1907a. The Cephalopoda from the deposit at the North End of False Bay, 

Zululand. Rep. geol. Surv. Natal Zululand, Pietermaritzburg, 3 : 163-234, pis 10-15. 

1907b. The Cephalopoda from the tributaries of the Manuan Creek, Zululand. Rep. 

geol. Surv. Natal Zululand, Pietermaritzburg, 3 : 235-249. 

Drushchitza, V. V. & Kudryavtseva, M. P. i960. Atlas of Lower Cretaceous fauna of the 
northern Caucasus and Crimea. 396 pp, 149 pis. Gostoptekhizdat, Moscow. [In Russian.] 


Fabre, S. 1940. Le cretace superieur de la Basse-Provence occidentale, i. Cenomanien et 

Turonien. Annls Fac. Sci. Marseille, Ser. II, 14 : 355 pp, 10 pis. 
Fallot, P. & Termier, H. 1923. Ammonites nouvelles des lies Baleares. Trah. Mus. 

nac. Cienc. nat. Madr., ser. Geol. 32 : 1-83, pis 1-6. 
Forbes, E. 1846. Report on the Cretaceous fossil invertebrates from southern India, collected 

by Mr. Kaye and Mr. CunUffe. Trans, geol. Soc. Lond., (2) 7 : 97-174, pis 7-19. 
Gabb, W. M. & Meek, F. B. 1864. Palaeontology of California, l:xx + 243 pp, 32 pis. 

Geol. Surv. Calif., San Francisco. 
Henderson, R. A. 1970. Ammonoidea from the Mata Series (Santonian-Maastrichtian) of 

New Zealand. Spec. Pap. Palaeont., London, 6 : 81 pp, 15 pis. 
Jones, D. L. 1963. Upper Cretaceous (Campanian and Maastrichtian) ammonites from 

southern Alaska. Prof. Pap. U.S. geol. Surv., Washington, 432 : 1-53, pis 1-41. 
Karakash, N. I. 1907. Le Cretace de la Crimee et sa faune. Trudy imp. S.-Peterb. Obshch. 

Estest., 32 : 1-482, pis 1-28. 
Kennedy, W. J. & Klinger, H. C. 1975. Cretaceous faunas from Zululand and Natal, 

South Africa. Introduction, Stratigraphy. Bull. Br. Mus. nat. Hist. (Geol.), London, 

25 (4) : 263-315, plate. 
Klinger, H. C, Wiedmann, J. & Kennedy, W. J. 1975. A new carinate phylloceratid 

ammonite from the early Albian (Cretaceous) of Zululand, South Africa. Palaeontology, 

London, 18 (3) : 657-664, pis 76-77. 
KossMAT, F. 1895-98. Untersuchungen iiber die sudindische Kreideformation. Beitr. 

Paldont. Geol. Ost-Ung., Vienna and Leipzig, 9 (1895) • 97-203 (1-107), pis 15-25 (i-ii) ; 

11 (1897)11-46 (108-153), pis 1-8 (12-19); 12 (1898) : 89-152 (154-217), pis 14-19 

McLearn, D. 1972. Ammonoids of the Lower Cretaceous sandstone member of the Haida 

Formation, Skidegate Inlet, Queen Charlotte Islands, western British Columbia. Bull. 

geol. Surv. Can., Ottawa, 188 : 1-78, 45 pis. 
Mahmoud, I. G. el-din 1955. Etudes paleontologiques sur la faune cretacique du massif du 

Moghara (Sinai, Egypte). Pubis Inst. Desert ^gypte, Heliopolis, 8 : 1-192, 19 pis. 
Marshall, P. 1926. The upper Cretaceous ammonites of New Zealand. Trans. Proc. N.Z. 

Inst., Wellington, 56 : 129-210, pis 19-47. 
Matsumoto, T. 1943. A short note on the Japanese Cretaceous Phylloceratidae. Proc. 

imp. Acad. Japan, Tokyo, 18 (for 1942) : 674-676. 

1953. The ontogeny of Metaplacenticeras subtilistriatum (Jimbo). Jap. J. Geol. Geogr., 

Tokyo, 23 : 139-150, pi. 13. 

1954- Appendix. Selected Cretaceous leading ammonites in Hokkaido and Saghalien. 

In Matsumoto, T. (Ed.) The Cretaceous System in the Japanese Islands : 243-313, pis 
17-36- Jap- Soc. Promotion Sci., Tokyo. 

1959- Cretaceous ammonites from the Upper Chitina Valley, Alaska. Mem. Fac. Sci. 

Kyushu Univ. (D), 8 : 49-90, pis 12-29. 

Michelin, H. 1834. Ammonites velledae. Magasin Zool., Paris, 3 : C IV, pi. 35. 
Neumayr, M. 1885. Die geographische Verbreitung der Juraformation. Denkschr. A had. 

Wiss. Wien, 50 : 57-142, pi. i. 
Oppel, a. 1865. Die Tithonische Etage. Z. dt. geol. Ges., Berlin, 17 : 535-558. 
Orbigny, a. d'. 1840-42. Paleontologie frangaise. Terrains Cretacds I. Cephalopodes. 

662 pp, 148 pis. Paris. 

1850. Prodrome de Paleontologie stratigraphique universelle des animaux mollusques et 

rayonnes, 2. 427 pp. Paris. 

Parona, C. F. & BoNARELLi, G. 1897. Fossih albani d'Escragnolles, del Nizzardo e della 
Liguria occidentale. Palaeontogr. ital., Pisa, 2 : 53-112, pis 10-14. 

Pervinquiere, L. 1907. Carte giologique de la Tunisie. Etudes de paleontologie tunisienne, 
I. Cephalopodes des terrains secondaires. 438 pp, 27 pis. Paris. 

1910. Sur quelques ammonites du Cretace algerien. Mem. Soc. gdol. Fr. PalSont., 

Paris, 42 : 1-86, 7 pis. 


Rawson, p. F. 1966. A phylloceratid ammonite from the Speeton Clay (Lower Cretaceous) 

of Yorkshire. Palaeontology, London, 9 : 455-457, pi. 72. 
Rousseau, L. 1842. Description des principaux fossiles de la Crimee. In Demidoff, A. de 

1840-42. Voyage dans la Russie meridionale et la Crimee par la Hongrie, la Valachie et la 

Moldavie (&c.) 2 : 781-823, Atlas pis 1-12. Paris. 
Sayn, G. 1 89 1. Description des ammonites nouvelles ou peu connues du Cretace inferieur 

du sud-est de la France. Mem. Soc. geol. Fr., Paris, 17 : 679-688, pi. 17. 

1901-07. Les ammonites pyriteuses des marnes valanginiennes du sud-est de la France. 

Mem. Soc. geol. Fr., Paris, 23 : 1-66, pis 1-6. 

1920. Les Phylloceras gargasiens du sud-est de la France (especes nouvelles ou peu 

connues). In Kilian, W. Contribution a I'etude des cephalopodes paleocretaces du 
sud-est de la France. Mem. Serv. Carte geol. det. France, Paris : 191-203, pi. i. 

Shimizu, S. 1935. The Upper Cretaceous cephalopods of Japan, Part I. /. Shanghai Sci. 

Inst., sect. 2, 1 : 159-226. 
SiMONOViCH, S., Batsevich, L. & SoROKiN, A. 1877. Description of the geology of the 

Kutais region etc. Mater. Geol. Kavk., Tifiis, 1876 : 1-112, pis 1-8. 
Spath, L. F. 1921a. On Cretaceous Cephalopoda from Zululand. Ann. S. Afr. Mus., 

Cape Town, 12 : 217-321, pis 19-26. 

1921b. On Upper Cretaceous Ammonoidea from Pondoland. Ann. Durban Mus., 

3 : 39-57. pis 6, 7. 

1922. On the Senonian ammonite fauna of Pondoland. Trans. R. Soc. S. Afr., Cape 

Town, 10 : 113-147, pis 5-9. 

— — 1923-43. A monograph of the Ammonoidea of the Gault. Palaeontogr. Soc. (Monogr.), 
London, 787 pp, 72 pis. 

1925. On Upper Albian Ammonoidea from Portuguese East Africa, with an appendix 

on Upper Cretaceous ammonites from Maputoland. Ann. Transv. Mus., Pretoria, 11 : 
179-200, pis 28-37. 

1927. Revision of the Jurassic Cephalopod fauna of Kachh (Cutch). Mem. geol. Surv. 

India Palaeont. indica, Calcutta, n.s. 11 : 1-71, pis 1-7. 

Stoliczka, F. 1863-66. The fossil Cephalopoda of the Cretaceous rocks of southern India. 

Mem. geol. Surv. India Palaeont. indica, Calcutta (3) 1 : 41-56, pis 26-31 (1863) ; 2-5 : 

57-106, pis 32-54 (1864) ; 6-9 : 107-154, pis 55-80 (1865) ; 10-13 : 155-216, pis 81-94 

Tate, R. 1865. Correlation of the Cretaceous Formations of the north-east of Ireland. 

Q. J I geol. Soc. Lond., 21 : 15-44, pls 3-6. 
ToucAS, A. 1890. Etude de la faune des couches Tithonique de I'Ardeche. Bull. Soc. geol. 

Fr., Paris, 18 : 560-630, pis 13-18. 
Usher, J. L. 1952. Ammonite faunas of the Upper Cretaceous of Vancouver Island, British 

Columbia. Bull. geol. Surv. Can., Ottawa, 21 : 1-182, pis 1-30. 
VAN HoEPEN, E. C. N. 1920. Description of some Cretaceous ammonites from Pondoland. 

Ann. Transv. Mus., Pretoria, 7 : 142-147, pis 24-26. 

1921. Cretaceous Cephalopoda from Pondoland. Ibid., 8 : 1-48, pis i-ii. 

Venzo, S. 1936. Cefalopodi del Cretacea medio-superiore dello Zululand. Palaeontogr. 

ital., Pisa, 36 : 59-133, pis 5-12. 
WiEDMANN, J. 1962a. Die systematische Stellung von Hypophylloceras Salfeld. Neues Jb. 

Geol. Paldont. Abh., Stuttgart, 115 : 243-256, pi. 16. 

1962b. Ammoniten aus der Vascogotischen Kreide (Nordspanien) . i. Phylloceratina, 

Lytoceratina. Palaeontographica, Stuttgart, A 118 ; 119-237, pis 8-14. 

1964. Unterkreide-Ammoniten von Mallorca, 2. Lieferung : Phylloceratina. Abh. 

math.-naturw. Kl. Akad. Wiss. Mainz, Jahr. 1963 (4) : 157-264, pis 11-21. 

1972. Neue Vorstellungen iiber Stammesgeschichte und System der Kreideammoniten. 

Proc. int. palaeont. Un., int. geol. Congr. XXIII Sess., Czechoslovakia ig68 : 93-120, 
I pi. Warsaw. 



WiEDMANN, J. & DiENi, I. 1968. Die Krcide Sardinians und ihre Cephalopoden. Palaeontogr. 

ital., Pisa, 64 : 1-171, 18 pis. 
Woods, H. 1906. The Cretaceous fauna of Pondoland. Ann. S. Afr. Mus., Cape Town, 

4 : 275-350, pis 33-44- 
YoKOYAMA, M. 1890. Versteinerungen aus der japanischen Kreide. Palaeontogr aphica, 

Stuttgart, 36 : 159-202, pis 18-25. 
ZiTTEL, K. A. 1868. Die Cephalopoden der Stramberger Schichten. In Oppel, A. Palaeont. 

Mitth. Mus. k. Bayer. St. 2 (i) : 1-118, pis 1-24. Stuttgart. 
ZwiERZYCKi, J. 1914. Die Cephalopodenfauna der Tendaguru-Schichten in Deutsch- 

Ostafrika. Arch. Biontol., Berlin, 3 (4) : 7-96, pis i-io. 


The page numbers of the principal references are printed in bold type. An asterisk (*) denotes 
a figure. 

acknowledgements 374 
Ammonites baborensis 371 

cylindricus 352 

partschi 369 

ramosus 359 

rouyanus 371 

serus 352 

subalpinus 355 

velledae 355, 360, 364 

zetes 352 
Aphroditiceras 354 ; see Salfeldiella 

Calaiceras 369 
Carinophylloceras 372-3 

collignoni 350, 372, 372*, 373-4, 374* ; pis 

Damesites 372-3 
? sp. nov. 373 
dimensions of specimens 351 

Epiphylloceras 351, 354, 363, 369, 371 

mikobokense 368 

radiatum 369 
erratum 380 
Euphylloceras 354 

serum 352 

velledae 360 

field locahties 351 

Geyeroceras see Phylloceras 
Goretophylloceras 354 

Hantkeniceras 351 
Harpophylloceras 372 

Heterophylloceras 351 
Hoplophylloceras 369 
Hypophylloceras 369 &c. ; see Phylloceras 

aphrodite 358 

cottreaui 369 

seresitense 364 

surya 369 
Hyporbulites 354, 363 

androiavense 365 

betieri 352, 354 

hoepeni 366-8, 368* 

seresitensis 364 
renaudiensis 364 

woodsi 366 

location of specimens 350 

Macrophylloceras 369 
Menegheniceras 372 

Neophylloceras 351, 354, 363 
hetonaiense 368 
lambertense 368 
marshalli 368 
nera 368 
seresitense 364 

Paraphylloceras 354 
Partschiceras 369 
baborense 371-2 
sp. juv. cf. baborense 350, 371-2 ; pi. 6, 

figs 2-3 
besairiei besairiei 370 

tsianalokensis 370 
bistriatum 370 



ezoense 370 

forbesianum 370-1 

inflatum 370 

? japonicum 356, 358 

marshalli 370 

minimum 370 

occlusum 370 

umzambiense 350, 370-1, 371* ; pi 

fig- 2 
whiteavesi 370 
zelandicum 370 
Partschiphylloceras 369 
Phylloceras 351 ; see also Hyporbulites 
androiavensis 365 
aphrodite 355, 358 
ellipticum 355-6, 358 
escragnollensis 358 
aff. grothi 358 
heterophyllum 351 
improvisum 356 
japonicum 356 
wera 366, 368 ; pi. 13, fig. 3 
onoense 352, 354 
cf. perlobatum 352 
ex aff. ramosi 364 
robionense 352-4 
sablyense 354 
semistriatum 352 
seresitense tanit 364-5 
serum 352, 354 

perlobata 352-3 
spathi 352-3 
aff. surya 368 
^Ae^:>/5 352, 354, 359 
umzambiense 370 
velledae 360-1, 364-5 

inflatum 362 

seresitense 364 
woodsi 366 
sp. 360-1, 366 
(Gey^rocems) 352 

cylindricus 352 
[Hypophylloceras) 352, 354-69 

algeriense 366 

androiavense 350, 365-6 ; pi. 4, fig. 7 

aphrodite 350, 355, 358-9 ; pi. 3, fig. 3 

aptiense applanatum 355, 358-9 
aptiense 355, 358-9 

aff. californicum 363 

cypris cypris 359, 362, 366, 372 
cytherae 359, 362, 366 

fortunei 355, 358 

hoepeni 368 

infundibuliformis 368 

improvisum 355-6, 358 
mikobokense 350, 368-9 ; pi. 12, fig. i 
mo^e^i 355, 358 
Me>'a 368 
onoense 352, 359 
pervinquierei 360 
ponticuli 359 
13, sablyense 355 

sardoum 355, 358 

seresitense 354, 359, 362, 363, 365 ; 
pi. 4, fig. 6 
boulei 363, 365 
seresitense 350, 363, 364-5 ; pi. 6, 

fig. 4 ; pi. 7, fig. 4 ; pi. 9 
/am7 363, 365 
subalpinum ellipticum 350, 354, 355-8, 
356*, 357*. 368 ; pi. 4, figs 1-4 ; 
pi. 5, figs 2-3 ; pi. 12, fig. 2 
subalpinum 354, 357-8, 368 
subseresitense 355, 358 
thetys diegoi 355, 358 
majoricense 355, 358 
/Ag/y5 354-5. 358-9 
velledae 354, 359, 360, 362, 372 ; pi. 6, 
fig. I 
aschiltaense 359-60, 362 
inflatum 350, 359-60, 362-3 ; pi. 13, 

fig. I 
morelianum 359-60, 362, 372 
velledae 350, 359, 360-2, 363, 372 ; 
pi. 4, fig. 5 ; pi. 5, fig. 4 ; pi. 7, 
figs 2-3 ; pis 8, 10, II 
sp. aff. velledae 350, 363 ; pi. 5, fig. 1 ; 

pi. 7, fig. I 
woodsi 367-8 

infundibuliformis 368 
woodsi 350, 366-8, 366*, 367*, 368* ; 
pi. 13, figs 3-5 
[Phylloceras] 351, 352-4 
heterophyllum 352 
serum 350, 352-4, 353* ; pis 1-2 ; pi. 3, 

figs 1-2 
5^a/;?i 352, 354 
[Salfeldiella] guettardi 358 
(Zetoceras) 352 
2-6^65 352 

Phyllocerataceae 351-74 
Phylloceratidae 351-74 
Phylloceratinae 351-74 
Phyllopachyceras 369 

umzambiense 370 
Procliviceras 369 
Pseudophylloceras 351 

Rhacoceras 351 

Salfeldiella [Aphroditiceras] see Phylloceras 
aphrodite venus 358 


Texanites oliveti Zone 368, 371 
Xeniophylloceras 351 
Zetoceras see Phylloceras 


The following error has been noted in the first paper of this series [Bull. Br. Mus. nat. Hist. 
(GeoL), 25 (4) : 263-315) : 

For 'Besaire' read 'Besairie' throughout. 

W. J. Kennedy 

Dept of Geology 6- Mineralogy 

University of Oxford 

Parks Road 

Oxford OXi 3PR 


H. C. Klinger 

South African Museum 
P.O. Box 61 
Cape Town 8000 
Republic of South Africa 

Accepted for publication 12 February 1976 


Phylloceras (Phylloceras) serum (Oppel) 

BM(NH) C78658, Makatini Formation, Barremian I, Loc. 170, Mlambongwenya Spruit, 

northern Znluland. x i. (p. 352 ; see also PI. 2 and PL 3, figs 1-2.) 

Bull. Br. Mus. nat. Hist. (Geol.) 27, 5 




Phylloceras (Phylloceras) serum (Oppel) 
Makatini Formation, Barremian I, Loc. 170, Mlambongwenya Spruit, northern Zululand. 

(p. 352 ; see also PI. i and PI. 3, figs 1-2.) 
Fig. la, b. BM(NH) C78660, x i. 
Fig. 2a-g. BM(NH) C78659. a-d, x i. e-g, x 2. 

Bull. Br. Mus. nat. Hist. (Geol.) 27, 5 



Phylloceras (Phylloceras) serum (Oppel) 
Makatini Formation, Barremian I, Loc. 170, Mlambongwenya Spruit, northern Zululand. 

X I. (p. 352 ; see also Pis 1-2.) 
Fig. la-c. BM(NH) C78657. 
Fig. 2a-b. BM(NH) C78661. 

Phylloceras (Hypophylloceras) aphrodite Fallot & Termier 

Fig. 3. BM(NH) C78691, Mzinene Formation, Albian V, Loc. 51 (Bed i), Mzinene River, 

Zululand. x i. (p. 358.) 

Bull. Br. Mils. nat. Hist. (Geol.) 27, 5 



Phylloceras (Hypophylloceras) subalpinum ellipticum Kossmat 
(P- 555 '> see also PL 5, figs 2-3 and PL 12, fig. 2.) All x i. 
Fig. I. BM(NH) C78665, Mzinene Formation, Albian V, Loc. 66, Munywana Creek 

Fig. 2. BM(NH) C78664, Mzinene Formation, Albian V, Loc. 54, Munywana Creek, Zululand. 
Fig. 3. BM(NH) C18139, figured Crick (1907a : pi. 10, figs 12, 12a), Mzinene Formation, 
Skoenberg region on the Lower Mzinene River, Zululand, probably Cenomanian III. 

Fig. 4. SAS A116, Mzinene Formation, Albian V, Loc. 67, Munywana Creek, Zululand. 

Phylloceras (Hypophylloceras) velledae velledae (Michelin) 

Fig. 5. BM(NH) C78683, Mzinene Formation, Albian V, Loc. 51 (Bed 6), Mzinene River, 
Zululand. x i. (p. 360 ; see also PL 5, fig. 4 ; PL 6, fig. i ; PL 7, figs 2-3 and Pis 8, 10, 11.) 

Phylloceras (Hypophylloceras) seresitense Pervinquiere 
Fig. 6. BM(NH) C18138, figured Crick (1907a : pi. 10, figs lo-ioa), Mzinene Formation, 
Cenomanian, Skoenberg region, Mzinene River, Zululand. x i. (p. 364; see also PL 6, 
fig. 4 ; PL 7, fig. 4 and PL 9.) 

Phylloceras (Hypophylloceras) androiavense Besairie 
Fig. 7. BM(NH) C78667, Mzinene Formation, Albian V, Loc. 54, Mzinene River, Zululand. 
X I- (P- 3^55-) 

Bull. Br. Mus. nat. Hist. (Geol.) 27, 5 



„ Phylloceras (Hypophylloceras) sp. aff. velledae (Michelin) 
Fig. I. BM(NH) C78696, Mzinene Formation, Albian V, Loc. 51, Mzinene River, Zululand 
(found loose). x i. (p. 363 ; see also PI. 7, fig. i.) 

Phylloceras (Hypophylloceras) subalpinum ellipticum Kossmat 
(P- 355 ; see also PL 4, figs 1-4 and PL 12, fig. 2.) All x 1-25. 
Fig. 2. BM(NH) C78663, Mzinene Formation, Albian V, Loc. 64 (Bed 4), Munywana Creek, 

Fig. 3. BM(NH) C78662, Mzinene Formation, Albian V, Loc. 51 (Bed 6), Mzinene River, 

Phylloceras (Hypophylloceras) velledae velledae (Michelin) 
Fig. 4. BM(NLi) C78672, Mzinene Formation, Albian V, Loc. 56, Mzinene River, Zululand. 
X I. (p. 360 ; see also PL 4, fig. 5 ; PL 6, fig. i ; PL 7, figs 2-3 and Pis 8, 10, 11.) 

Bull. Br. Mus. nat. Hist. (Geol.) 27, 5 



Phylloceras (Hypophylloceras) velledae (Michelin) 
Fig. I. SAS Ziga, from Haughton's (1936:291, fig. 3) Loc. 19. x i. (p. 360; typical 
form see also PL 4, fig. 5 ; PI. 5, fig. 4 ; PL 7, figs 2-3 and Pis 8, 10, 11.) 

Partschiceras sp. juv. ? cf. baborense (Coquand) 
Figs 2-3. BM(NH) C78693, Makatini Formation, Barremian I, Loc. 170, Mlambongwenya 
Spruit, northern Zululand. x i ; 3a-3b x 2. (p. 371.) 

Phylloceras (Hypophylloceras) seresitense seresitense Pervinquiere 
Fig. 4. BM(NH) C18137, figured Crick (1907a : pi. 10, figs ii-iia), Mzinene Formation, 
Cenomanian, Skoenberg Region, Mzinene River, Zululand. x i. (p. 364 ; see also PL 4, fig. 6 ; 
PL 7, fig. 4 and PL 9.) 

Bull. Br. Mus. nat. Hist. (Geol.) 27, 5 



■"> *.r' II 

.'. ^ :''>r 


Phylloceras (Hypophylloceras) sp. aff. velledae (Michelin) 
Fig. I. B]\I(NH) C78695, Mzinene Formation, Albian V, Loc. 56, Mzinene River, Zululand. 
XI. (p. 363 ; see also PI. 5, fig. i.) 

Phylloceras (Hypophylloceras) velledae velledae (Michelin) 
(p. 360; see also PI. 4, fig. 5 ; PI. 5, fig. 4 ; PL 6, fig. i and Pis 8, 10, 11.) Both x i. 
Fig. 2. BM(NH) C78673, Mzinene Formation, Albian V, Loc. 56, Mzinene River, Zululand. 
Fig. 3. BM(NH) C78684, Mzinene Formation, Albian V, Loc. 51 (Bed 12 or 13), Mzinene 
River, Zululand. 

Phylloceras (Hypophylloceras) seresitense seresitense Pervinquiere 
Fig. 4. BM(NH) C78690, Mzinene Formation, ? Cenomanian III, The Skoenberg, Mzinene 
River, Zululand. x i. (p. 364 ; see also PI. 4, fig. 6 ; PI. 6, fig. 4 and PL g.) 

Bull. Br. Mus. nat. Hist. (Geol.) 27, 5 


/■#- •" 


Phylloceras (Hypophylloceras) velledae velledae Michelin 

BM(NH) C78668, Mzinene Formation, Albian IV, Loc. 51 (Bed i), Mzinene River, Zululand. 

X 0-9 ; actual diameter 208 mm. (p. 360 ; see also PI. 4, fig. 5 ; PL 5, fig. 4 ; PI. 6, fig. i ; 

PI. 7, figs 2-3 and Pis 10- 11.) 

Bull. Br. Mus. nat. Hist. (Geol.) 27, 5 




Phylloceras (Hypophylloceras) seresitense seresitense Pervinquiere 

SAS A1401, Mzinene Formation, Albian III, Loc. 36, Mzinene River, Zululand. 

X 074 ; actual diameter 190 mm. (p. 364 ; see also PL 4, fig. 6 ; PL 6, fig. 4 and PL 7, fig. 4.) 

Bull. Br. Mus. nat. Hist. (Geol.) 27, 5 


^ ^ . 

C ■-- V' 

^ - 



Phylloceras (Hypophylloceras) velledae velledae (Michelin) 

SAS Zi492a, Mzinene Formation, Albian V, Loc. 54, Mzinene River, Zululand. 

X 0-86 ; actual diameter 162 mm. (p. 360 ; see also PL 4, fig. 5 ; PI. 5, fig. 4 ; 

PL 6, fig. I ; PL 7, figs 2-3 and Pis 8, 11.) 

Bull. By. Mus. nat. Hist. (Geol.) 27, 5 


., V 


"■-i.s» ■/.- 


Phylloceras (Hypophylloceras) velledae velledae (Michelin) 
SAS Z1496, Mzinene Formation, Albian V, Mzinene River, Zululand. 
X 0-85 ; actual diameter 169 mm. (p. 360 ; see also PL 4, fig. 5 ; PI. 5, fig. 
PL 6, fig. I ; PL 7, figs 2-3 and Pis 8, 10.) 

Bull. Br. Mus. nat. Hist. (Geol.) 27, 5 



^ - ' ' ^i. '._ '\ ' ^^• ^.^ •-• . ~¥t-.r' '^-■»| V'J 


Phylloceras (Hypophylloceras) mikobokense (Collignon) 
Fig. I. SASH163C/7, St Lucia Formation, Maastrichtian I, Loc. 20, Mfolozi River, 
Zululand. x i. (p. 368.) 

Phylloceras (Hypophylloceras) subalpinum ellipticum Kossmat 
Fig. 2. BM(NH) C78666, Mzinene Formation, Cenomanian II or III, The Skoenberg, 
Mzinene River, Zululand. x i. (p. 355 ; see also PI. 4, figs 1-4 and PI. 5, figs 2-3.) 

Bull. Br. Mus. nat. Hist. (Geol.) 27. 5 






-»■ ml 



Phylloceras (Hypophylloceras) velledae inflatum Collignon 
Fig. I. SASZ1742, Mzinene Formation, Albian IV or V, Mzinene River, Zululand. x i. 

(p. 362.) 

Partschiceras umzambiense (van Hoepen) 
Fig. 2. Holotype, TM 524, Umzamba Formation, Santonian or early Campanian, Loc. i, 
mouth of Umzamba River, southern Natal (Pondoland). x i. (p. 370.) 

Phylloceras (Hypophylloceras) woodsi woodsi van Hoepen 
Umzamba Formation, Santonian or early Campanian, Loc. i, 
mouth of Umzamba River, southern Natal (Pondoland). (p. 366.) 
Fig. 3. Durban Museum specimen, mentioned by Spath (1921 : 50) as Phylloceras nera 
(Forbes). x i. 

Fig. 4. Paratype, TM 534, x 2. 
P~iG. 5. Holotype, TM 537, x 2. 

Bull. Br. Mus. nat. Hist. (Geol.) 27, 5 



*•■■,'' l* 

r- % 





"^ '*■• 

M ^^ 



ft p 

'^ % 




Carinophylloceras coUignoni Klinger, Wiedmann & Kennedy 

Paratype, BM(NH) C78640, Mzinene Formation, Albian III, Loc. 36, Mzinene River, 

Zululand. x i. (p. 373 ; see also PL 15.) 

Bull. Br. Mus. nat. Hist. (Geol.) 27, 5 



Carinophylloceras collignoni Klinger, Wiedmann & Kennedy 
Albian III, Mzinene River, Zululand. Both x i. (p. 373 ; see also PL 14. 
Fig. I. Paratype, BM(NH) C78644, Loc. 35 (Bed 2). 
Fig. 2. Paratype, BM(NH) C78768, Loc. 36. 

Bull. Br. Mus. nat. Hist. (Cxeol.) 27, 5 


-^ ■ • ■ L 

'Mi '^ '% 

^-. 4> 



/ \ 






1. Cox, L. R. Jurassic Bivalvia and Gastropoda from Tanganyika and Kenya. 
Pp. 213 ; 30 Plates ; 2 Text-figures. 1965. out of print. 

2. El-Naggar, Z. R. Stratigraphy and Planktonic Foraminifera of the Upper 
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Egypt, U.A.R. Pp. 291 ; 23 Plates ; 18 Text-figures. 1966. ;^ii. 

3. Davey, R. J., DowNiE, C, Sarjeant, W. a. S. & Williams, G. L. Studies on 
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figures. 1966. £8,20. 

3. Appendix. Davey, R. J., Downie, C, Sarjeant, W. A. S. & Williams, G. L. 
Appendix to Studies on Mesozoic and Cainozoic Dinoflagellate Cysts. Pp. 24. 
1969. 95p. 

4. Elliott, G. F. Permian to Palaeocene Calcareous Algae (Dasycladaceae) of 
the Middle East. Pp. iii ; 24 Plates ; 16 Text-figures. 1968. out of print. 

5. Rhodes, F. H. T., Austin, R. L. & Druce, E. C. British Avonian (Carboni- 
ferous) Conodont faunas, and their value in local and continental correlation. 
Pp. 313 ; 31 Plates ; 92 Text-figures. 1969. £13.10. 

6. Childs, a. Upper Jurassic RhynchonelUd Brachiopods from Northwestern 
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7. Goody, P. C. The relationships of certain Upper Cretaceous Teleosts with 
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8. Owen, H. G. Middle Albian Stratigraphy in the Anglo-Paris Basin, Pp. 164 ; 
3 Plates ; 52 Text-figures. 1971. ^^7.20. 

9. SiDDiQui, Q. A. Early Tertiary Ostracoda of the family Trachyleberididae 
from West Pakistan. Pp. 98 ; 42 Plates ; 7 Text-figures. 1971. £9.60. 

10. FoREY, P. L. a revision of the elopiform fishes, fossil and Recent. Pp. 222 ; 
92 Text-figures. 1973. £ii.35- 

11. Williams, A. Ordovician Brachiopoda from the Shelve District, Shropshire. 
Pp. 163 ; 28 Plates ; 11 Text-figures ; no Tables. 1974. £12.80. 

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